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The Urban Book Series
Ransford A. Acheampong
Spatial
Planning in
Ghana
Origins, Contemporary Reforms and
Practices, and New Perspectives
Series Advisory Editors

Fatemeh Farnaz Arefian, University College London, London, UK
Michael Batty, University College London, London, UK
Simin Davoudi, Newcastle University, Newcastle, UK
Geoffrey DeVerteuil, Cardiff University, Cardiff, UK
Karl Kropf, Oxford Brookes University, Oxford, UK
Karen Lucas, University of Leeds, Leeds, UK
Marco Maretto, University of Parma, Parma, Italy
Fabian Neuhaus, University of Calgary, Calgary, Canada
Vítor Oliveira, Porto University, Porto, Portugal
Christopher Silver, University of Florida, Gainesville, USA
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Igor Vojnovic, Michigan State University, East Lansing, USA
Jeremy Whitehand, University of Birmingham, Birmingham, UK
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Spatial Planning in Ghana

Origins, Contemporary Reforms
and Practices, and New Perspectives
123
Department of Planning and Environmental
Management, School of Environment,
Education and Development
The University of Manchester
Manchester, UK
ISSN 2365-757X ISSN 2365-7588 (electronic)

ISBN 978-3-030-02010-1 ISBN 978-3-030-02011-8 (eBook)
https://doi.org/10.1007/978-3-030-02011-8
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My mother, Florence Adwoa Sekyiwaa
Hackman
Preface
In 2012, I had the unique opportunity to be involved in a new wave of institutional

and legal reforms that would reintroduce and establish a new spatial planning
system in Ghana. These reforms had actually begun in 2007, while I was still an
undergraduate student of planning, under the Land Use Planning and Management
Project (LUPMP). LUPMP, one of the sub-component initiatives under the
umbrella of the Land Administration Project (LAP), was an ambitious attempt to
resurrect a hitherto weak and under-resourced apparatus for land use planning and
management in Ghana. Having freshly graduated from a master’s degree pro-
gramme in 2012, I had the privilege of working in one of the teams of consultants
that had been tasked, to among other things, provide technical assistance in the
preparation of the very first set of spatial development plans. Our task was quite
significant in that it would help experiment with the various spatial planning models
initiated by the reforms and pave the way to institutionalize the new spatial plan-
ning system.
For myself and for spatial planning—a discipline and profession that I love and
have been passionate about—this was a historic moment. For the first time, since
the first national physical development plan for Ghana was formulated around
1963, spatial planning was receiving the much needed attention in both the public
policy and academic discourse. This was certainly a great impetus for the profes-
sional practice of spatial planning and the roles it could play in addressing some
of the key developmental challenges of the country. Students of spatial planning,
including myself, beamed with hope and optimism not only about the career
opportunities a well-resourced spatial planning system could provide, but also about
the opportunity this would afford us to put our skills to use to address some of the
challenges facing our towns and cities.
I really enjoyed the experience of working with my team. I also received
practical knowledge and invaluable insights from working closely and interacting
with practitioners in the public sector (i.e. Town Planning Officers) who had spent
decades doing what they possibly could to salvage a profession that was essentially
at the brink of demise. That said, on the scholarly side of things, I found a major
gap. Historically, spatial planning as a discipline has attracted limited scholarship in
vii
viii Preface
Ghana. This may partly be as a result of the historically weak land use planning
system and the limited attention that the profession has received in public policy
discourse in the past. Yet another reason may be the shift in the focus and content of
planning education in the early 1990s, which emphasized the social and economic
dimensions of planning and development to the neglect of their physical expression
and distributional consequences within and between geographical entities (i.e.
neighbourhoods, towns, cities and regions).
The motivation to write this book, the first of its kind on the subject in Ghana,
therefore, stems partly from my passion for spatial planning as a profession and the
practical knowledge and first-hand experience I obtained through my involvement
in the delivery of some of the initiatives under LUPMP, which would contribute to
the establishment of a new spatial planning system in Ghana. I have always
believed that one of the most effective ways by which the future cadre of planners
would learn from these significant developments that have shaped spatial planning
would be if we can document the experiences and lessons while reflecting on ways
of improving the system and the associated practices to deliver its objectives.
Towards this end, I conceived the idea to write this book.
Furthermore, this book is the result of my desire as a researcher to contribute to
the scholarship needed to expand the frontiers of knowledge in the discipline while
searching for ways of making spatial planning useful in addressing the country’s
developmental challenges. Indeed, traditionally, books on spatial planning have
been written from Global North perspectives. In my student days, both in Ghana
and abroad, the key textbooks I read on the topic were mostly written from either
European or North American experiences. These sources have offered the oppor-
tunity to engage with the experiences and practices from their respective contexts.
However, we know that the experience in one context may not necessarily be
relevant in other contexts. Similarly, what has been proven to work in one context
might not necessarily work in another. This is particularly true for spatial planning
scholarship and practice because planning education and planning systems tend to
be deeply rooted in the sociocultural, legal, political and economic situations of the
countries of origin. I see this book on spatial planning in Ghana as being a part of an
ongoing effort to advance scholarship and knowledge on the subject from contexts
that have received limited attention in the past.
By focusing on Ghana, the book aims to provide an in-depth and comprehensive
discussion of the spatial planning system and associated practices from this context.
It hopes to take its audience on a journey to discover and understand spatial
planning from a unique social, cultural, legal and political-economy context.
Readers will engage with the wide-ranging issues covering the systems, processes,
procedures, mechanisms and practices that characterize the spatial planning system
and shape outcomes in this context. The book has been written with undergraduate
and graduate students, academic researchers and practitioners and policy-makers in
the multidisciplinary field of spatial planning in mind. In particular, this book aims
at readers who seek an international perspective in spatial planning systems and
practices.
Preface ix
Many individuals have helped me in cultivating the knowledge, skills and the
motivation that would enable me to write this book. I am particularly thankful to my
teachers for the inspiring lectures I have received on the subject. They passed on
useful knowledge and nurtured in me the passion that would ultimately culminate in
writing this book. I must thank Prince Aboagye Anokye, my long-time mentor from
my undergraduate days at the Kwame Nkrumah University of Science and
Technology. On his recommendation, I got the opportunity to get closer to the action
and to experience first-hand, some of the major initiatives that would help establish
the new spatial planning system in Ghana—an experience that would also be
indispensable in writing this book. I also want to thank Priscilla Ankomah-Hackman
for helping me in the digital production of most of the maps in this book.
I am grateful for the support I have received from the editorial team at Springer,
especially from Juliana Pintanguy, Selvaraj Divya, Sanjievkumar Mathiyazhagan
and Komala Jaishankar towards the publication of this book.
Finally, I could not have completed this book without the support of my wife,
Nana Adwoa Nyarko, who kept the distractions to the barest minimum and also
proofread most of the initial draft chapters of this book. What will I do without your
unconditional support and encouragement, Nana?
Dublin, Ireland Ransford A. Acheampong, Ph.D.

July 2018
Acknowledgements
I would like to thank Stephen Asabere, Priscilla Ankomah-Hackman and

Mohammed Abdul-Fatawu for providing, free of charge, the built-up land-cover
datasets used for the analyses of urbanization trends in Ghana. Stephen provided the
datasets on Accra; Priscilla provided the datasets on the Sekondi-Takoradi
Metropolis, and Mohammed provided the datasets covering the Greater Kumasi
sub-region.
Some of the chapters in this book build on the evidence from initial works I have
done and/or co-published with other colleagues. Notable among these works are:
(i) ‘Housing for the urban poor: towards alternative financing strategies for
low-income housing development in Ghana’ (with P. A. Anokye) published in
International Development Planning Review (37(4), 445–465, 2015); (ii) ‘One
nation, two planning systems? Spatial planning and multi-level policy integration in
Ghana: Mechanisms, challenges and the way forward’ (with Ibrahim Alhassan)
published in Urban Forum (27 (1) 1–18, 2016); (iii) ‘Quantifying the spatio-temporal
patterns of settlement growth in a metropolitan region of Ghana’ (with Felix
S. K. Agyemang and Mohammed Abdul-Fatawu) published in GeoJournal (82(4),
823–840, 2017); and (iv) ‘Examining the determinants of utility bicycling using a
socio-ecological framework: An exploratory study from the Tamale Metropolis in
Northern Ghana’ (with Siiba Alhassan) published in the Journal of Transport
Geography (article in press, 2018). I would like to thank my co-authors for allowing
me to build on some of the ideas and information initially presented in these works.
I would also like to thank Mr Kofi Amedzro Kekeli (TCPD Head Office, Accra)
for helping me to obtain the necessary permission from the TCPD to use some
of the diagrams and illustrations in this book. Finally, I want to thank the two
anonymous individuals who reviewed and provided feedback on the book proposal.
I am particularly grateful for their very useful comments and suggestions and,
above all, for their encouragement which further motivated me to write this book.
xi
Contents
Part I Setting the Scene

1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
2 The Concept of Spatial Planning and the Planning System . . . . . . . 11
2.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
2.2 What Is Spatial Planning? . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
2.3 Spatial Planning and Political-Economy Contexts . . . . . . . . . . . 15
2.3.1 State-Market Interactions: Market Failure,
the Welfare State and Spatial Planning . . . . . . . ..... 15
2.4 Normative Theories of Spatial Planning: An Overview . . ..... 19
2.5 The Spatial Planning System: Key Features and Policy
Instruments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..... 21
2.5.1 Governance Structure and Spatial Planning . . . . ..... 22
2.5.2 Policy Instruments Deployed to Achieve Spatial
Planning Objectives . . . . . . . . . . . . . . . . . . . . . ..... 22
2.6 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..... 25
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..... 26
3 Historical Origins and Evolution of Spatial Planning
and the Planning System in Ghana . . . . . . . . . . . . . . . . . . . . . . . .. 29
3.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 29
3.2 Origins of Spatial Planning in Ghana . . . . . . . . . . . . . . . . . . .. 30
3.2.1 Pre-1945 Colonial Planning: Essentially Spatial Plans
in Character . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 31
3.2.2 The 1945 Town and Country Planning
Act (CAP 84) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 32
xiii
xiv Contents
3.3 Landmark (Spatial) Plan-Making Projects in the Era

of Independence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 34
3.3.1 National Perspectives—Five-Year Plan (1951–1956)
and Seven-Year Development Plan
(1963/64–1969/70) . . . . . . . . . . . . . . . . . . . . . . . . . .. 35
3.3.2 The National Physical Development Plan
(1963–1970) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
3.3.3 The Master Plan for Accra . . . . . . . . . . . . . . . . . . . . . 37
3.3.4 The Master Plan for Tema . . . . . . . . . . . . . . . . . . . . . 41
3.3.5 Planning Scheme for Kumasi, 1963 . . . . . . . . . . . . . . . 43
3.4 Decentralization and Contemporary Planning in Ghana . . . . . . . 45
3.5 The Age of Reform: The Land Use Planning and Management
Project and the New Spatial Planning System . . . . . . . . . . . . .. 48
3.5.1 The Land Use and Spatial Planning Act (Act 925),
2016 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 49
3.5.2 Institutional Framework for Spatial Planning Under
the Land Use and Spatial Planning Act (Act 925) . . .. 50
3.5.3 The Three-Tier, Hierarchical System of Spatial
Planning Instruments . . . . . . . . . . . . . . . . . . . . . . . .. 51
3.5.4 The Land Use and Spatial Planning Development
Fund . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 52
3.6 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 53
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 56
Part II Modern-Day Planning Systems and Scales

of Spatial Planning
4 Contemporary Traditions of Planning and Spatial Planning
at the National Level . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 59
4.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 60
4.2 Three Decades of National Medium-Term Development
Planning . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 60
4.3 The Systems Act and the New Land Use and Spatial Planning
Act: A Recipe for Institutional Conflicts and Duplications? . . .. 65
4.4 National-Level Spatial Planning Under the New Spatial
Planning System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 68
4.4.1 The Ghana National Spatial Development Framework
(2015–2035) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 69
4.4.2 The National Urban Policy Framework . . . . . . . . . . .. 76
4.5 Realizing Long-Term Spatial Development Imperatives
Within a Culture of Medium-Term Development Planning . . .. 77
4.6 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 79
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 81
Contents xv
5 The Inception of Regional Spatial Planning . . . . . . . . . . . . . . . . .. 83

5.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 83
5.2 The Concept of Regions and Rationale for Regional Spatial
Planning . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 85
5.3 Spatial Planning for Regions in Ghana: A Previously
Uncharted Territory . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 88
5.4 Contemporary Planning Systems and Authority
and Competences for Regional Spatial Planning . . . . . . . . . . .. 90
5.5 Regional Spatial Planning in Practice Under the New Spatial
Planning System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 93
5.5.1 Western Region Spatial Development Framework
(2012–2022) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 93
5.5.2 Greater Kumasi Sub-Regional Spatial Development
Framework (2013–2033) . . . . . . . . . . . . . . . . . . . . . .. 95
5.6 The Emerging Paradigm of Special Development Zones
and Development Authorities . . . . . . . . . . . . . . . . . . . . . . . . . . 99
5.7 Towards Effective Regional Spatial Planning . . . . . . . . . . . . . . 100
5.8 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
6 Local-Level Spatial Planning and Development Management . . . . . 107
6.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108
6.2 Local-Level Spatial Planning Instruments . . . . . . . . . . . . . . . . . 109
6.2.1 District Spatial Development Framework . . . . . . . . . . . 109
6.2.2 Structure Plan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
6.2.3 Local Plan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110
6.3 The Context of Local Spatial Planning in Ghana . . . . . . . . . . . 111
6.3.1 Four Decades of Severed Connection Between
National Policy Frameworks and Local Land Use
Plans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111
6.3.2 A Complex Land Tenure System and the Separation
Between Ownership Rights and Determination
of Use of Land . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113
6.3.3 Legal and Institutional Arrangements
for Contemporary Local Spatial Planning
and Development Management . . . . . . . . . . . . . . . . . . 114
6.4 Local Plans and Development Management in Practice . . . . . . . 118
6.4.1 Plan-Led Development versus Non-Plan
Development-First and the Permitting Process . . . . . . . 119
6.4.2 Rezoning and Change of Use . . . . . . . . . . . . . . . . . . . 126
xvi Contents
6.5 Non-Compliance with Local Plans and Development

Management Regulations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129
6.6 Towards Effective Local Spatial Planning and Development
Management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134
Part III Issues in Spatial Planning

7 Policy Integration in Spatial Planning: Mechanisms,
Practices and Challenges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
7.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
7.2 The Concept of Integration in Spatial Planning . . . . . . . . . . . . . 140
7.2.1 Types and Dimensions of Policy Integration . . . . . . . . 141
7.3 Integration in Ghana’s Spatial Planning System:
Mechanisms, Practices and Challenges . . . . . . . . . . . . . . . . . . . 142
7.4 Towards Effective Policy Integration in Spatial Planning . . . . . . 147
7.5 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
8 Public Engagement in Spatial Planning: Statutory Requirements,
Practices and Challenges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151
8.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151
8.2 Public Participation in Theory . . . . . . . . . . . . . . . . . . . . . . . . . 152
8.3 Spatial Planning and Public Participation in Ghana: Practices
and Challenges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 156
8.3.1 Public Participation Requirements and Practices
Under the 1945 Town and Country Planning
Ordinance (CAP 84) . . . . . . . . . . . . . . . . . . . . . . . . . . 156
8.3.2 Democratization, Decentralization and Public
Participation in Spatial Planning: Just More
of the Same? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157
8.3.3 Public Participation Under the New Spatial Planning
System: Practices and Emerging Issues . . . . . . . . . . . . 159
8.4 Towards Effective Stakeholder Engagement: The Case
for Technology-Mediated Participation Practices . . . . . . . . . . . . 166
8.5 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168
9 Urbanization and Settlement Growth Management . . . . . . . . . . . . . 171
9.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171
9.2 Global Urbanization Trends and the Processes of Urban
Change . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173
9.3 Urbanization Trends in Ghana . . . . . . . . . . . . . . . . . . . . . . . . . 176
9.3.1 Urban Population Growth . . . . . . . . . . . . . . . . . . . . . . 177
9.3.2 Spatio-temporal Settlement Expansion Trends . . . . . . . 179
Contents xvii
9.4 Urbanization and Urban Growth Management . . . . . . . . . . . . . 192

9.4.1 Regulatory Instruments . . . . . . . . . . . . . . . . . . . . . . . . 193
9.4.2 Fiscal Instruments—Taxes, Exactions and Fees . . . . . . 197
9.4.3 Incentive-based Instruments . . . . . . . . . . . . . . . . . . . . 198
9.5 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 199
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201
10 The Spatial Planning System and Housing Development:
Prospects and Models . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205
10.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205
10.2 The Housing Situation in Ghana . . . . . . . . . . . . . . . . . . . . . . . 207
10.2.1 Channels of Housing Supply and Supply-Side
Challenges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 208
10.2.2 Demand-Side Challenges . . . . . . . . . . . . . . . . . . . . . . 211
10.3 Policy Responses in the Past . . . . . . . . . . . . . . . . . . . . . . . . . . 215
10.4 The Planning System and Housing Delivery: Models
and Approaches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
10.4.1 Model 1: Supplying Land for New Housing
Development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
10.4.2 Model 2: Development Exactions and Affordable
Housing Quotas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222
10.5 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227
11 Integrated Spatial Development and Transportation Planning . . . . 231
11.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231
11.2 Conceptualizing the Spatial Development and Transportation
Nexus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233
11.2.1 Spatial Development and Transportation
at the Strategic Level . . . . . . . . . . . . . . . . . . . . . . . . . 235
11.2.2 Spatial Development and Transportation at the City
and Neighbourhood Levels . . . . . . . . . . . . . . . . . . . . . 238
11.3 Urban Spatial Structure and Mobility Patterns in Ghana . . . . . . 241
11.3.1 A Conceptual Model of the Structure of Metropolitan
Regions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 242
11.3.2 Exemplifying the Generalized Model of Urban
Spatial Structure the Case of the Greater Kumasi
Metropolitan Region . . . . . . . . . . . . . . . . . . . . . . . . . . 245
11.3.3 The Influence of Prevailing Spatial Structures
on Commuting Patterns and Travel Behaviours . . . . . . 249
xviii Contents
11.4 Towards Integrated Spatial Development and Transport Policy

and Planning . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251
11.4.1 Land Use and Transport Integration at Policy
and Institutional Levels . . . . . . . . . . . . . . . . . . . . . . . . 252
11.4.2 Urban Planning and Development to Promote
Sustainable, Non-Motorized Mobility . . . . . . . . . . . . . 258
11.4.3 A Portfolio of Complementary Policy Strategies:
Public Transit, Shared-Mobility and Intermodal
Integration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 261
11.4.4 The Role of Integrated Land Use and Transport
Decision-Support Systems . . . . . . . . . . . . . . . . . . . . . . 263
11.5 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 264
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 265
12 Spatial Planning and the Urban Informal Economy . . . . . . . . . . . . 269
12.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 269
12.2 Informal Urbanism and the Urban Informal Economy:
An Overview of Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . 271
12.3 Historical Origins and Contemporary Manifestations
of the Informal Economy . . . . . . . . . . . . . . . . . . . . . . . . . . . . 273
12.4 Location Decisions and Challenges of Urban Informal
Economy Businesses and Workers . . . . . . . . . . . . . . . . . . . . . . 275
12.4.1 Home-Based Informal Economic Activities . . . . . . . . . 276
12.4.2 Itinerant Informal Economy Workers . . . . . . . . . . . . . . 277
12.4.3 Sedentary Sales/Commercial and Artisanal Activities
Located Outside the Home . . . . . . . . . . . . . . . . . . . . . 278
12.5 Planning to Accommodate the Informal Economy . . . . . . . . . . . 280
12.5.1 Shifting Attitudes and Contemporary Policy
Responses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281
12.5.2 Translating National Policies into Local Strategies
and Actions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 283
12.6 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 286
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 287
13 Epilogue: Perspectives on Pathways Towards a Responsive
Spatial Planning System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 289
13.1 A Synopsis of the Story so Far . . . . . . . . . . . . . . . . . . . . . . . . 289
13.2 Strengthening the Spatial Planning System . . . . . . . . . . . . . . . . 291
13.3 Responding to Developmental Challenges Through Spatial
Planning . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 294
13.3.1 Local-Level Spatial Planning and Development
Control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 294
13.3.2 Urbanization Impact Monitoring and Settlement
Growth Management . . . . . . . . . . . . . . . . . . . . . . . . . 296
Contents xix
13.3.3 Housing Supply—The Role of Spatial Planning . . . . . . 297

13.3.4 Integrated Spatial Development and Transport
Planning for Sustainable Mobility . . . . . . . . . . . . . . . . 298
13.3.5 Supporting Livelihoods and Local Economic
Development—Spatial Planning and the Urban
Informal Economy . . . . . . . . . . . . . . . . . . . . . . . . . . . 299
13.3.6 Public Engagement for the Co-production of Urban
Transformations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 300
13.4 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 301
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 302
About the Author
Ransford A. Acheampong Ph.D., M.Phil. Cambridge, UK; B.Sc. KNUST, Ghana

is a Presidential Academic Fellow at the University of Manchester, UK. Before
joining the University of Manchester, he was a Postdoctoral Research Fellow at
Trinity College Dublin, The University of Dublin, Ireland. His research focuses on
the nexus between spatial development and transportation. Themes covered in his
research include comparative study of spatial planning systems; land use
and transport interaction modelling; socio-spatial impacts of emerging mobility
paradigms (e.g. shared-mobility and autonomous transport); and travel behaviours,
focusing on the socio-ecological determinants of sustainable mobility options such
as bicycling and walking in cities. He has contributed to planning-related projects
for national and international organizations, including as a spatial planning con-
sultant for the Town and Country Planning Department (TCPD) in Ghana and a
research consultant for the Organization for Economic Co-operation and
Development (OECD).
xxi
Abbreviations
AMA Accra Metropolitan Area/Assembly

AUN Aflao Urban Network
BAU Business Advisory Unit
BRT Bus Rapid Transit
CBD Central Business District
CCUN Cape Coast Urban Network
CEDECOM Central Region Development Commission
CSGs Civil society groups
DACF District Assemblies Common Fund
DEPP Development Policy and Planning
DPA District Planning Authorities
DPU Development Planning Unit
DVLA Driver and Vehicle Licencing Authority
ESPON European Spatial Planning Observation Network
GDP Gross Domestic Product
GKSR Greater Kumasi Sub-Region
GKSR-SDF Greater Kumasi Sub-Regional Spatial Development
Framework
GLSS Ghana Living Standards Survey
GPRS Growth and Poverty Reduction Strategy
GREDA Ghana Real Estate Developers Association
GSGDA Ghana Shared Growth and Development Agenda
HHUN Ho-Hohoe Urban Network
HIPC Heavily Indebted Poor Countries
ICT Information and Communication Technology
IDA International Development Association
ILO International Labour Organization
IMF International Monetary Fund
IPEP Infrastructure for Poverty Eradication Project
JICA Japan International Cooperation Agency
xxiii
xxiv Abbreviations
KMA Kumasi Metropolitan Area/Assembly

KNUST Kwame Nkrumah University of Science and Technology
LAP Land Administration Project
LP Local Plan
LUPMP Land Use Planning and Management Project
LUSPA Land Use and Spatial Planning Authority
LUTI Land Use Transport Interaction
MDGs Millennium Development Goals
MESTI Ministry of Environment, Science, Technology and Innovation
METLOMP-SIM Metropolitan Location and Mobility Patterns Simulator
MLGRD Ministry of Local Government and Rural Development
MMDA Municipal, Metropolitan and District Assembly
MTDP Medium-Term Development Plan
NCA The National Communications Authority
NDC National Democratic Congress
NDPC National Development Planning Commission
NEUN North-East Urban Network
NGOs Non-Governmental Organizations
NHP National Housing Policy
NIP National Infrastructure Plan
NPF National Policy Framework
NPP New Patriotic Party
NRSC National Road Safety Commission
NSDF National Spatial Development Framework
NSPS National Spatial Planning System
NTP National Telecommunications Policy
NTP National Transport Policy
NUP National Urban Policy
OECD Organization for Economic Co-operation and Development
OfD Oil for Development
PIP Public Investment Programmes
RCC Regional Co-ordinating Council
RPCU Regional Planning Co-ordinating Unit
RSD Regional Spatial Development Framework
RSPC Regional Spatial Planning Committee
SADA Savannah Accelerated Development Authority
SAP Structural Adjustment Programme
SDF Spatial Development Framework
SDGs Sustainable Development Goals
SP Structure Plan
SPC Statutory Planning Committee
SPRING Spatial Planning for Regions in Growing Economies
STMA Sekondi-Takoradi Metropolitan Area/Assembly
STUN Sekondi-Takoradi Urban Network
SUN Sunyani Urban Network
Abbreviations xxv
TAZs Traffic Analysis Zones

TCDP Town and Country Planning Department
TDR Transfer of Development Rights
TOD Transit-oriented development
TUN Tamale Urban Network
UBG Urban growth boundary
USB Urban service boundary
WRSDF Western Regional Spatial Development Framework
WUN Wa Urban Network
List of Figures
Fig. 2.1 Constituents of a typical spatial planning system . . . . . . . . . . . . 21

Fig. 3.1 Proposed residential densities in the 1958 Master Plan for
Accra. Source Reproduced digitally from analogue maps
obtained from the TCPD, Accra. . . . . . . . . . . . . . . . . . . . . . . . . 38
Fig. 3.2 Distribution of major open spaces in the 1958 Master Plan for
Fig. 3.3 Designated as slums and areas proposed as suitable for
lowering of construction standards in the 1958 Master Plan for
Fig. 3.4 A typical neighbourhood plan accompanying the 1958 Master
Plan for Accra for private development at Kaneshie, Accra.
Source 1958 Master Plan for Accra, TCPD, Accra . . . . . . . . . . 40
Fig. 3.5 Underlying principles of spatial organization in the Tema
Master Plans. Adapted from Doxiadis Associates (1962) . . . . . . 41
Fig. 3.6 Proposed road system in the Tema Master Plan. Source
Reproduced digitally from analogue map obtained from the
TCPD, Accra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
Fig. 3.7 Distribution of communities based on income groups in the
Tema Master Plan. Source Reproduced digitally from analogue
map obtained from TCPD, Accra . . . . . . . . . . . . . . . . . . . . . . . . 43
Fig. 3.8 Planning Scheme for Kumasi, 1963. Source TCPD, Kumasi . . . 44
Fig. 3.9 Governance structure and institutional competences under the
National Development Planning (System) Act (Act 480, 1994)
and the Land Use and Spatial Planning Act (Act, 925, 2016).
Source Acheampong and Alhassan (2016, p. 7) . . . . . . . . . . . . . 46
Fig. 3.10 Governance structure and institutional competences for spatial
planning under the 2016 Land Use and Spatial Planning Act
(Act 925) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
xxvii
xxviii List of Figures
Fig. 3.11 Three-tier model of spatial development frameworks and

derivative plans established by the new decentralized spatial
planning system . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Fig. 4.1 Institutional arrangements for planning under two main
planning laws in Ghana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
Fig. 4.2 Composition of the governing body of the Land Use and
Spatial Planning Authority (LUSPA) . . . . . . . . . . . . . . . . . . . . . 67
Fig. 4.3 Urbanization trends in Ghana. Source Based on Population
and Housing Census data, Ghana Statistical Services
(For more information see: http://www.statsghana.gov.gh/
statistics.html) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
Fig. 4.4 Regional population growth rate (1960–2010). Source Based
on Population and Housing Census data, Ghana Statistical
Services . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
Fig. 4.5 ‘Integrated spatial development concept’ proposed by the
NSDF. Source Ghana National Spatial Development
Framework . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
Fig. 5.1 Composition of the Regional Spatial Planning Committee
established by the Land Use and Spatial Planning
Act (Act 925) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
Fig. 5.2 Diagram of the Western Region spatial development
framework. Source Town and Country Planning Department,
Accra Head Office . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
Fig. 5.3 Diagram of the Greater Kumasi Sub-Regional Spatial
Development Framework. Source Town and Country Planning
Department, Kumasi Ghana . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
Fig. 6.1 Physical development and development control under
normative plan-led versus non-plan scenarios . . . . . . . . . . . . . . 121
Fig. 8.1 Arnstein’s (1969) ladder of participation . . . . . . . . . . . . . . . . . . 153
Fig. 8.2 Pretty’s typology of participation. Adapted
from Pretty (1995) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154
Fig. 9.1 Percentage of the population living in urban areas
(1950–2050). Source World urbanization prospects: based on
world urbanization prospects data, United Nations (2014) . . . . . 173
Fig. 9.2 Built-up area expansion for a globally representative sample of
200 cities (1990–2014). Source Based on data obtained
from Atlas of urban expansion, 2016 edition (Atlas report can be
downloaded at: http://www.lincolninst.edu/sites/default/files/
pubfiles/atlas-of-urban-expansion-2016-volume-1-full.pdf.
Raw data files were downloaded at website: http://www.
atlasofurbanexpansion.org/data) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175
Fig. 9.3 National population growth trend (1950–2050). Source Based
on world urbanization prospects data,
United Nations (2014) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177
List of Figures xxix
Fig. 9.4 Percentage of urban population in Ghana, West Africa and the
African Continent (1950–2050). Source Based on world
urbanization prospects data, United Nations (2014) . . . . . . . . . . 178
Fig. 9.5 Urban population growth rate in Ghana, West Africa and the
African Continent (1950–2050). Source Based on world
urbanization prospects data, United Nations (2014) . . . . . . . . . . 179
Fig. 9.6 Spatio-temporal built-up land-cover changes in the Accra
metropolis and surrounding areas . . . . . . . . . . . . . . . . . . . . . . . . 182
Fig. 9.7 Spatio-temporal built-up land-cover changes in the Greater
Kumasi sub-region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187
Fig. 9.8 Spatio-temporal built-up land-cover changes in the
Sekondi-Takoradi Metropolis . . . . . . . . . . . . . . . . . . . . . . . . . . . 190
Fig. 11.1 A three-level conceptual illustration of the links between
spatial development and transportation . . . . . . . . . . . . . . . . . . . . 234
Fig. 11.2 Road and railway transport infrastructure networks
in Ghana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 237
Fig. 11.3 Fibre-optic networks for ICT in Ghana . . . . . . . . . . . . . . . . . . . 239
Fig. 11.4 A conceptual illustration of the spatial structure of
metropolitan regions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 242
Fig. 11.5 Map of the GKSR showing local government
administrative areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 246
Fig. 11.6 Three broad urban zones in the KMA . . . . . . . . . . . . . . . . . . . . 247
Fig. 11.7 Distribution of broad land use activity functions
in the KMA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 248
Fig. 11.8 A conceptual illustration major land use activity functions
in the KMA and road distances between them . . . . . . . . . . . . . . 249
Fig. 11.9 Institutional setting for integration between spatial
development and transportation policy and planning . . . . . . . . . 255
List of Tables
Table 3.1 Periodizing significant developments in the history of spatial

planning in Ghana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
Table 4.1 Spatial planning-related issues captured in national policy
frameworks under the tradition of development planning . . . . 63
Table 4.2 Historical population size and growth rate . . . . . . . . . . . . . . . . 70
Table 4.3 Historical population distribution in the ten administrative
regions of Ghana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
Table 4.4 Proposed hierarchy of settlement in the NSDF . . . . . . . . . . . . 73
Table 4.5 City-regions and urban networks identified and delineated
in the NSDF . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
Table 5.1 Proposed hierarchy of settlement in the WRSDF. . . . . . . . . . . 95
Table 5.2 Spatial Development Zones . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Table 9.1 Historical population growth in major towns and cities
in Ghana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179
Table 9.2 Land-cover changes in Ghana between 1975 and 2013 . . . . . . 181
Table 9.3 Built-up land changes in Accra and surround districts
between 1984 and 2015 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183
Table 9.4 Aggregated built-up land changes in Accra and surrounded
districts between 1984 and 2015 . . . . . . . . . . . . . . . . . . . . . . . 184
Table 9.5 Built-up land changes in Greater Kumasi sub-region between
1986 and 2014 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 186
Table 9.6 Population distribution in the Greater Kumasi sub-region . . . . 186
Table 9.7 Built-up land changes in Sekondi-Takoradi between 1990
and 2016 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190
Table 9.8 Types of growth management instruments . . . . . . . . . . . . . . . . 193
Table 10.1 Annual household incomes housing expenditure ratio . . . . . . . 212
xxxi
List of Boxes
Box 6.1 Functions of the District Spatial Planning Committee and

Technical Subcommittee under the Land Use and Spatial
Planning Act (Act 925, 2016) . . . . . . . . . . . . . . . . . . . . . . . . .. 116
Box 6.2 Composition of District Spatial Planning Committee
and Technical Subcommittee under the Land Use
and Spatial Planning Act (Act 925, 2016) . . . . . . . . . . . . . . . .. 116
Box 6.3 Development Zones in the Zoning Guidelines
and Planning Standards . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 122
Box 6.4 Conditions for Change of Use . . . . . . . . . . . . . . . . . . . . . . . . .. 127
Box 6.5 Procedure for Processing Rezoning and Change of Use
Applications . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 128
Box 8.1 Stages and Methods of Consultation in SDF
and Local Plan Preparation . . . . . . . . . . . . . . . . . . . . . . . . . . .. 160
Box 9.1 The Greater Kumasi Urban Growth Boundary . . . . . . . . . . . . .. 195
Box 10.1 Examples of Properties Supplied in the Private Real-Estate
Market in Accra, Ghana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 214
Box 10.2 Housing Policy Statements in Ghana . . . . . . . . . . . . . . . . . . . .. 216
Box 11.1 Ministries and Agencies in the Transportation Sector . . . . . . .. 252
Box 11.2 The National Transport Policy: Sector Goals
and Strategies for Land Use and Transport Integration . . . . . .. 254
Box 11.3 National Transport Policy: Policy Statement
and Strategies for Non-motorized Transport . . . . . . . . . . . . . . .. 260
Box 12.1 Informal Business Zone (BL) . . . . . . . . . . . . . . . . . . . . . . . . . .. 282
xxxiii
Part I
Setting the Scene
Chapter 1
Introduction
Abstract This chapter provides a general introduction for the book. It sets out the
aim, scope and structure of this book. It provides a brief account of the story of
spatial planning in Ghana, highlighting landmark events in spatial planning from
the mid-1940s when the planning system was first established to the contemporary
times. Following from this, it introduces the three main parts of the book and provides
an overview of the themes in spatial planning that are subsequently explored in the
remaining twelve chapters of the book.
Keywords Spatial planning · Institutions · Legal systems · Socio-economic

Development · Global south · Ghana
Planning systems are deeply rooted in the sociocultural, legal, political and economic
situations of the countries that use them to achieve specific development goals. In
view of this, scholarship on the subject tends to take a country-specific focus and/or
a comparative approach by addressing spatial planning issues and experiences from
multiple countries. Historically, scholarship on spatial planning has based their nar-
ratives on perspectives and experiences from the Global North. The Global North
perspectives and experiences have therefore not only made significant contribution
to our understanding of the evolving meaning, scope, purpose and practices of spa-
tial planning, but also shaped spatial planning theory development and the models
that are currently applied globally. In recent years, however, there have been calls to
explore, analyse and document knowledge and experiences from different contexts
with respect to spatial planning. This, in turn, has generated a huge interest in and
demand for authoritative perspectives on contemporary spatial planning from other
contexts, particularly in countries in the Global South.
Efforts are currently underway to respond to the gaps in the existing academic lit-
erature by documenting spatial planning practices and experiences from previously
unexplored contexts. While some of these initial attempts have focused exclusively
on spatial planning as it relates to urban issues in multiple countries, others written as
compendiums, due to their relatively wider scope, have understandably been unable
to offer detailed discussions on the wider context of national governance structures
and political-economy ethos that shape the spatial planning systems, practices and
© Springer Nature Switzerland AG 2019 3

R. A. Acheampong, Spatial Planning in Ghana, The Urban Book Series,
https://doi.org/10.1007/978-3-030-02011-8_1
4 1 Introduction
outcomes within the individual countries they cover.1 Thus, in addition to the afore-
mentioned endeavours, it is vital that we continue to have scholarly engagements with
spatial planning from unique sociocultural, legal and political-economy contexts on
wide-ranging issues, including the systems, processes, procedures, mechanisms and
practices that characterize the spatial planning system and shape outcomes in par-
ticular national contexts. This book is intended to be one of the responses to this
imperative.
This book is about spatial planning in Ghana. Although the main focus is on
Ghana, it situates the discussion within the wider contemporary discourse on the
evolving meaning, scope and purpose of spatial planning as both an established
academic discipline and a public-sector activity for addressing a plethora of social,
economic and environmental challenges. The mandate and expected outcomes of
spatial planning may be similar across different contexts. Yet, the actual scope of
issues embraced by the activity could be wide-ranging, depending on the prevailing
sociopolitical, economic, legal and cultural systems within which the activity takes
place and from which it derives its mandate and relevance.
The aim of this book is not to address every possible issue that may be delineated
and addressed through spatial planning. Instead, the core of this book engages with
some of the dominant themes in spatial planning globally, with a particular focus on
detailed analyses of the experiences and practices from the unique context of Ghana.
It provides a comprehensive and critical discussion on the changing socio-economic
conditions, institutional settings, legal arrangements and practices which have shaped
and defined Ghana’s spatial planning system for more than seven decades. The book
traces the history of spatial planning in Ghana to identify the landmark initiatives
that have been responsible for introducing, reforming and institutionalizing spatial
planning as we know it today. One of the major themes addressed in this book
is the geographical scale of spatial planning, where we examine the role of the
spatial planning system in formulating programmes, policies and plans at the national,
regional and local levels of governance and decision-making. The issue of policy
integration across spatial scales and between policy domains in spatial planning is
also addressed in this book. Adopting a problem-based approach and drawing on
primary and secondary data sources, the book also engages with important themes
in the field, including: public participation in spatial planning; urbanization and
sustainable growth management; spatial planning and housing development; spatial
planning and the transition towards sustainable urban transport; and spatial planning
and the informal economy.
Spatial planning in Ghana has had a chequered history. In the first decade fol-
lowing independence from British rule, physical planning, as it was then called,
became one of the most important machineries for articulating and realizing the
ambitious programme of socio-economic transformation envisioned by the socialist
government of Dr. Kwame Nkrumah. Over this period, landmark spatial plan-making
projects including the formulation of the first National Physical Development Plan
1 See, e.g., Silva (2015) and Bhan et al. (2018).

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as a Natica or Buccinum perforates the shell of a Tellina or Mactra in
order to get at its contents.
Few observations appear to have been made on the habits or
food of Streptaxis, Rhytida, Ennea, Daudebardia, Paryphanta, and
other carnivorous Mollusca. A specimen of Ennea sulcata, enclosed
in the same box as a Madagascar Helix (sepulchralis Fér.) many
times its own size, completely emptied the shell of its inhabitant.[126]
Mr. E. L. Layard informs me that certain Cape Rhytida, e.g. R.
capsula Bens., R. dumeticola Bens., and R. vernicosa Kr., eat
Cyclostoma affine, Helix capensis, H. cotyledonis, etc. To Mr. Layard
I am also indebted for the—perhaps apocryphal—tradition that the
best time to capture the great Aerope caffra Fér. in numbers was
after an engagement between the Kaffirs and Zulus, when they
might be observed streaming from all points of the compass towards
the field of slaughter. The Cuban Oleacina are known to secrete a
very bitter fluid which they emit; this perhaps produces a poisoning
or benumbing effect upon their victims when seized. They devour
operculates, e.g. Helicina regina and sagraiana.[127]
CHAPTER III
ENEMIES OF THE MOLLUSCA—MEANS OF DEFENCE—MIMICRY AND
PROTECTIVE COLORATION—PARASITIC MOLLUSCA—COMMENSALISM—
VARIATION
Enemies of the Mollusca

The juicy flesh and defenceless condition of many of the Mollusca
make them the favourite food and often the easy prey of a host of
enemies besides man. Gulls are especially partial to bivalves, and
may be noticed, in our large sandy bays at the recess of the tide,
busily devouring Tellina, Mactra, Mya, Syndosmya, and Solen. On
the Irish coast near Drogheda a herring gull has been observed[128]
to take a large mussel, fly up with it in the air over some shingly
ground and let it fall. On alighting and finding that the shell was
unbroken it again took it up and repeated the process a number of
times, flying higher and higher with it until the shell was broken.
Hooded crows, after many unavailing attempts to break open
mussels with their beak, have been seen to behave in a similar way.
[129] Crows, vultures, and aquatic birds carry thousands of mussels,
etc., up to the top of the mountains above Cape Town, where their
empty shells lie in enormous heaps about the cliffs.[130]
The common limpet is the favourite food of the oyster-catcher,
whose strong bill, with its flattened end, is admirably calculated to
dislodge the limpet from its seat on the rock. When the limpet is
young, the bird swallows shell and all, and it has been calculated
that a single flock of oyster-catchers, frequenting a small Scotch
loch, must consume hundreds of thousands of limpets in the course
of a single year. Rats are exceedingly fond of limpets, whose shells
are frequently found in heaps at the mouth of rat holes, especially
where a cliff shelves gradually towards a rocky shore. A rat jerks the
limpet off with a sudden movement of his powerful jaw, and, judging
from the size of the empty shells about the holes, has no difficulty in
dislodging the largest specimens. ‘I once landed,’ relates a shepherd
to Mr. W. Anderson Smith,[131] ‘on the I. of Dunstaffnage to cut
grass, and it was so full of rats that I was afraid to go on; and the
grass was so full of limpets that I could scarcely use the scythe, and
had to keep sharpening it all the time.’ Sometimes, however, the
limpet gets the better both of bird and beast. The same writer
mentions the case of a rat being caught by the lip by a limpet shell,
which it was trying to dislodge. A workman once observed[132] a bird
on Plymouth breakwater fluttering in rather an extraordinary manner,
and, on going to the spot, found that a ring dotterel had somehow
got its toe under a limpet, which, in closing instantly to the rock, held
it fast. Similar cases of the capture of ducks by powerful bivalves are
not uncommon, and it is said that on some parts of the American
coasts, where clams abound, it is impossible to keep ducks at all,
[133] for they are sure to be caught by the molluscs and drowned by
the rising tide.
The Weekly Bulletin of San Francisco, 17th May 1893, contains
an account of the trapping of a coyote, or prairie wolf, at Punta
Banda, San Diego Co., by a Haliotis Cracherodii. The coyote had
evidently been hunting for a fish breakfast, and finding the Haliotis
partially clinging to the rock, had inserted his muzzle underneath to
detach it, when the Haliotis instantly closed down upon him and kept
him fast prisoner.
Rats devour the ponderous Uniones of North America. When Unio
moves, the foot projects half an inch or more beyond the valves. If,
when in this condition, the valves are tightly pinched, the foot is
caught, and if the pinching is continued the animal becomes
paralysed and unable to make use of the adductor muscles, and
consequently flies open even if the pressure is relaxed. The musk-rat
(Fiber zibethicus) seizes the Unio in his jaws, and by the time he
reaches his hole, the Unio is ready to gape.[134] Rats also eat
Vivipara, and even Limnaea, in every part of the world.
Every kind of slug and snail is eaten greedily by blackbirds,
thrushes, chaffinches, and in fact by many species of birds. A thrush
will very often have a special sacrificial stone, on which he dashes
the shells of Helix aspersa and nemoralis, holding them by the lip
with his beak, until the upper whorls are broken; heaps of empty
shells will be found lying about the place of slaughter. The bearded
Titmouse (Parus biarmicus) consumes quantities of Succinea putris
and small Pupa, which are swallowed whole and become triturated
in the bird’s stomach by the aid of numerous angular fragments of
quartz.[135]
Frogs and toads are very partial to land Mollusca. A garden
attached to the Laboratory of Agricultural Chemistry at Rouen had
been abandoned for three years to weeds and slugs. The director
introduced 100 toads and 90 frogs, and in less than a month all the
slugs were destroyed, and all kinds of vegetables and flowers,
whose cultivation had until then been impossible, were enabled to
flourish.[136]
Certain Coleoptera are known to prey upon Helices and other
land Mollusca. Récluz noticed, near Agde, a beetle (Staphylinus
olens) attack Helix ericetorum when crawling among herbage,
sticking its sharp mandibles into its head. Every time the snail
retreated into its shell the beetle waited patiently for its
reappearance, until at last the snail succumbed to the repeated
assaults. M. Lucas noticed, at Oran, the larva of a Drilus attacking a
Cyclostoma. The Drilus stood sentinel at the mouth of a shell, which
was closed by the operculum, until the animal began to issue forth.
The Drilus then with its mandibles cut the muscle which attaches the
operculum to the foot, disabling it sufficiently to prevent its being
securely closed, upon which it entered and took possession of the
body of its defenceless host, completing its metamorphosis inside
the shell, after a period of six weeks.[137] The female glow-worm
(Lampyris noctiluca) attacks and kills Helix nemoralis.
Among the Clavicornia, some species of Silpha carry on a
determined warfare against small Helices. They seize the shell in
their mandibles, and then, throwing their head backwards, break the
shell by striking it against their prothorax.
The common water beetle, Dytiscus marginalis, from its strength
and savage disposition, is a dangerous enemy to fresh-water
Mollusca. One Dytiscus, kept in an aquarium, has been noticed to kill
and devour seven Limnaea stagnalis in the course of one afternoon.
The beetles also eat L. peregra, but apparently prefer stagnalis, for
when equal quantities of both species were placed within their reach,
they fixed on the latter species first.[138]
In East Africa a species of Ichneumon (Herpestes fasciatus)
devours snails, lifting them up in its forepaws and dashing them
down upon some hard substance.[139] In certain islands off the south
coasts of Burmah, flat rocks covered with oysters are laid bare at low
tide. A species of Monkey (Macacus cynomolgus) has been noticed
to furnish himself with a stone, and knock the oysters open, always
breaking the hinge-end first, and then pulling out the mollusc with his
fingers.[140]
The walrus is said to support himself almost entirely on two
species of Mya (truncata and arenaria), digging them out of the
sand, in which they live buried at a depth of about 1½ feet, with his
powerful tusks. Whales swallow enormous numbers of pelagic
molluscs (Clio, Limacina), which are at times so abundant in the
Arctic seas, as to colour the surface for miles. Many of the larger
Cetacea subsist in great part on Cephalopoda; as many as 18 lbs. of
beaks of Teuthidae have been taken from the stomach of a single
Hyperoodon.
Fish are remarkably partial to Mollusca of various kinds. The cat-
fish (Chimaera) devours Pectunculus and Cyprina, crushing the stout
shells with its powerful jaws, while flounders and soles content
themselves with the smaller Tellina and Syndosmya which they
swallow whole. As many as from 30 to 40 specimens of Buccinum
undatum have been taken from the stomach of a single cod, and the
same ‘habitat’ has been recorded for some of the rarer whelks, e.g.
Bucc. humphreysianum, Fusus fenestratus, the latter also occurring
as the food of the haddock and the red gurnard. No less than 35,000
Turtonia minuta have been found in the stomach of a single mullet.
Nudibranchs are no doubt dainty morsels for fish, and hence have
developed, in many cases, special faculties for concealment, or, if
distasteful, special means of remaining conspicuous (see pp. 71–
74).
Fig. 22.—Two valves of Mytilus
edulis L., representing
diagrammatically the
approximate position of
the holes bored by
Purpura in about 100
specimens of Mytilus,
gathered at Newquay,
Cornwall.
Besides the dangers to which they are exposed from other
enemies, many of the weaker forms of Mollusca fall a prey to their
own brethren. Nassa and Murex on this side of the Atlantic, and
Urosalpinx on the other, are the determined foes of the oyster.
Purpura lapillus prefers Mytilus edulis to any other food, piercing the
shell in about two days’ time by its powerful radula, which it appears
to employ somewhat in gimlet fashion. If Mytilus cannot be procured,
it will eat Littorina or Trochus, but its attempts on the hard shell of
Patella are generally failures. The statement which is sometimes
made, that the Purpura makes its hole over the vital parts of the
Mytilus, appears, according to the evidence embodied in the
annexed figure, to be without foundation. The fact is that a hole in
any part of its shell is fatal to the Mytilus, since the long proboscis of
the Purpura, having once made an entrance, can reach from one
end of the shell to the other. The branchiae are first attacked, the
adductor muscles and edges of the mantle last. Natica and Nassa
pierce in a similar way the shells of Mactra, Tellina, Donax, and
Venus. Murex fortispina is furnished with a powerful tooth at the
lower part of its outer lip. At Nouméa, in New Caledonia, its favourite
food is Arca pilosa, which lives half buried in coral refuse. The Murex
has been seen to drag the Arca from its place of concealment, and
insert the tooth between the valves, so as to prevent their closing,
upon which it was enabled to devour its prey at leisure.[141]
The carnivorous land Mollusca, with the exception of Testacella,
appear to feed by preference upon other snails (pp. 54, 55).
Parasitic Worms, Mites, etc.—A considerable number of the
Trematode worms pass one or more of the stages in the cycle of
their development within the bodies of Mollusca, attaining to the
more perfect or sexual form on reaching the interior of some
vertebrate. Thus Distoma endolabum Duj. finds its first intermediate
host in Limnaea stagnalis and L. ovata, its second in L. stagnalis, or
in one of the fresh-water shrimps (Gammarus pulex), or in the larvae
of one of the Phryganeidae (Limnophilus rhombicus), attaining to the
sexual form in the common frog. Distoma ascidia v. Ben. passes
firstly through Limnaea stagnalis or Planorbis corneus, secondly
through certain flies and gnats (Ephemera, Perla, Chironomus), and
finally arrives within certain species of bats. Distoma nodulosum
Zed. inhabits firstly Paludina impura, secondly certain fishes
(Cyprinus Acerina), and lastly the common perch. The sporocyst of
Distoma macrostomum inhabits Succinea putris, pushing itself up
into the tentacles, which become unnaturally distended (Fig. 23).
While in this situation it is swallowed by various birds, such as the
thrush, wagtail, and blackbird, which are partial to Succinea, and
thus obtains lodgment in their bodies. Amphistoma subclavatum
spends an early stage in Planorbis contortus, after which it becomes
encysted on the skin of a frog. When the frog sheds its skin, it
swallows it, and with it the Amphistoma, which thus becomes
established in the frog’s stomach.[142]
Fig. 23.—A Trematode worm
(Leucochloridium paradoxum
Car.) parasitic in the tentacles
of Succinea putris L. × 20
(after Baudon).
The common liver-fluke, which in the winter of 1879–1880 cost
Great Britain the lives of no less than three million sheep, is perhaps
the best known of these remarkable parasitic forms of life. Its history
shows us, in one important particular, how essential it is for the
creature to meet, at certain stages of its existence, with the exact
host to which it is accustomed. Unless the newly-hatched embryo
finds a Limnaea truncatula within about eight hours it becomes
exhausted, sinks, and dies. It has been tried with all the other
common pond and river Mollusca, with Limnaea peregra, palustris,
auricularia, stagnalis, with Planorbis marginatus, carinatus, vortex,
and spirorbis, with Physa fontinalis, Bithynia tentaculata, Paludina
vivipara, as well as with Succinea putris, Limax agrestis and
maximus, Arion ater and hortensis. Not one of them would it touch,
except occasionally very young specimens of L. peregra, and in
these its development was arrested at an early stage. But on
touching a L. truncatula the embryo seems to know at once that it
has got what it wants, and sets to work immediately to bore its way
into the tissues of its involuntary host, making by preference for the
branchial chamber; those which enter the foot or other outlying parts
of the Limnaea proceed no farther.[143]
Many similar cases occur, in which littoral Mollusca, such as
Littorina and Buccinum, form the intermediate host to a worm which
eventually arrives within some sea-bird.
Certain Nematode worms (Rhabditis) are known to inhabit the
intestine of Arion, and the salivary glands of Limax agrestis. Diptera
habitually lay their eggs within the eggs of Helix and Limax. Many
species of mite (Acarina) infest land Pulmonata. No adult Limax
maximus is without at least one specimen of Philodromus (?)
limacum, and the same, or an allied species, appears to occur on the
larger of our Helices, retiring upon occasion into the pulmonary
chamber.
Several of the Crustacea live associated with certain molluscs.
Pinnotheres lives within the shell of Pinna, Ostrea, Astarte,
Pectunculus, and others. Apparently the females alone reside within
the shell of their host, while the males seize favourable opportunities
to visit them there. A specimen of the great pearl-oyster (Meleagrina
margaritifera) was recently observed which contained a male
Pinnotheres encysted in nacre. It was suggested that he had
intruded at an unfortunate time, when no female of his kind
happened to be in, and that, having penetrated too far beneath the
mantle in the ardour of his search, was made prisoner before he
could escape.[144] Ostracotheres Tridacnae lives in the branchiae of
the great Tridacna. A little brachyurous crustacean inhabits the raft of
Ianthina, and assumes the brilliant blue colour of the mollusc.
Means of Defence
As a rule, among the Mollusca, the shell forms a passive mode of
resistance to the attacks of enemies. Bivalves are enabled, by
closing their valves, to baffle the assault of their smaller foes, and
the operculum of univalves, both marine and land, serves a similar
purpose. Many land Mollusca, especially Helix and Pupa, as well as
a number of Auriculidae, have the inside of the aperture beset with
teeth, which are sometimes so numerous and so large that it is
puzzling to understand how the animal can ever come out of its
shell, or, having come out, can ever draw itself back again. Several
striking cases of these toothed apertures are given in Fig. 24.
Whatever may be the origin of these teeth, there can be little doubt
that their extreme development must have a protective result in
opposing a barrier to the entrance, predatory or simply inquisitive, of
beetles and other insects. Sometimes, it will be noticed (G), the
aperture itself is fairly simple, but a formidable array of obstacles is
encountered a little way in. It is possible that the froth emitted by
many land snails has a similar effect in involving an irritating intruder
in a mass of sticky slime. The mucus of slugs and snails, on the
other hand, is more probably, besides its use in facilitating
locomotion, a contrivance for checking evaporation, by surrounding
the exposed parts of their bodies with a viscid medium.
Fig. 24.—Illustrating the elaborate arrangement of teeth in the

aperture of some land Pulmonata. A. Helix (Labyrinthus)
bifurcata Desh., Equador. B. H. (Pleurodonta) picturata
Ad., Jamaica. C. H. (Dentellaria) nux denticulata Chem.,
Demerara. D. Anostoma carinatum Pfr., Brazil; a, tube
communicating with interior of shell. E. H. (Stenotrema)
stenotrema Fér., Tennessee, × 3/2. F. H. (Polygyra)
auriculata Say, Florida, × 3/2. G. H. (Plectopylis) refuga
Gld., Tenasserim (a and b × 2).
Some species of Lima shelter themselves in a nest constructed of
all kinds of marine refuse, held together by byssiferous threads.
Modiola adriatica, M. barbata, and sometimes M. modiolus conceal
themselves in a similar way. Gastrochaena frequently encloses itself
in a sort of half cocoon of cement-like material. The singular genus
Xenophora protects itself from observation by gluing stones, shells,
and various débris to the upper side of its whorls (Fig. 25).
Sometimes the selection is made with remarkable care; the
Challenger, for instance, obtained a specimen which had decorated
its body whorl exclusively with long and pointed shells (Fig. 26).
Fig. 25.—Xenophora (Phorus)

conchyliophora Born.,
concealed by the stones
which it glues to the upper
surface of its shell. (From a
British Museum specimen.)
Fig. 26.—Xenophora (Phorus)
pallidula Reeve. A mollusc
which escapes detection by
covering itself with dead
shells of other species. (From
a Challenger specimen in the
British Museum, × ½.)
The formidable spines with which the shells, e.g. of the Murex
family, are furnished must contribute greatly to their protection
against fishes, and other predatory animals. Murex tenuispina, for
instance (see chap. ix.), would prove as dangerous a morsel in the
mouth of a fish as a hedgehog in that of a dog. Whether the singular
tooth in the outer lip of Leucozonia (see chap. xiv.), a feature which
is repeated, to a less marked extent, in Monoceros and several of
the West Coast muricoids, is developed for defensive purposes,
cannot at present be decided.
The Strombidae possess the power of executing long leaps,
which they doubtless employ to escape from their foes. In their case
alone this power is combined with singular quickness of vision. On
one occasion Mr. Cuming, the celebrated collector, lost a beautiful
specimen of Terebellum, by the animal suddenly leaping into the
water, as he was holding and admiring it in his hand. Miss Saul has
informed me that the first living specimen of Trigonia that was ever
obtained was lost in a similar way. It was dredged by Mr. Stutchbury
in Sydney Harbour, and placed on the thwart of a small boat. He had
just remarked to a companion that it must be a Trigonia, and his
companion had laughed at the idea, reminding him that all known
Trigonia were fossil, when the shell in question baffled their efforts to
discover its generic position by suddenly leaping into the sea, and it
was three months before Mr. Stutchbury succeeded in obtaining
another.
Some genera possess more than merely passive means of
defence. Many Cephalopoda emit a cloud of inky fluid, which is of a
somewhat viscous nature, and perhaps, besides being a means of
covering retreat, serves to entangle or impede the pursuer. The
formidable suckers and hooks possessed by many genera in this
Order are most dangerous weapons, both for offence and defence.
Aplysia, when irritated, ejects a purple fluid which used to be
considered dangerously venomous. Many of the Aeolididae,
including our own common Aeolis papillosa, possess stinging cells at
the end of their dorsal papillae, the effect of which is probably to
render them exceedingly distasteful to fish.
The common Vitrina pellucida has a curious habit which in all
probability serves for a defence against birds in the winter. When
crawling on the edge of a stone or twig it has the power of suddenly
jerking its ‘tail,’ so as to throw itself on the ground, where it is
probably lost to sight among decaying leaves. At other times it rolls
away a few inches and repeats the jump. It also possesses the
power of attaching to itself bits of leaves or soil, which entirely cover
and conceal both shell and animal.[145] The property of parting with
the tail altogether, a remarkable form of self-defence, has already
been noticed on p. 44.
The poisonous nature of the bite of certain species of Conus is
well authenticated. Surgeon Hinde, R.N., saw[146] a native on the I.
of Matupi, New Britain, who had been bitten by a Conus geographus,
and who had at once cut small incisions with a sharp stone all over
his arm and shoulder. The blood flowed freely, and the native
explained that had he not taken these precautions he would have
died. Instances have been recorded of poisonous wounds being
inflicted by the bite of Conus aulicus, C. textile, and C. tulipa.
According to Mr. J. Macgillivray[147] C. textile at Aneitum (S. Pacific)
is called intrag, and the natives say it spits the poison upon them
from several inches off! Two cases of bites from C. textile occurred
to this gentleman’s notice, one of which terminated fatally by
gangrene. Sir Edward Belcher, when in command of the Samarang,
was bitten[148] by a Conus aulicus at a little island off Ternate in the
Moluccas. As he took the creature out of the water, it suddenly
exserted its proboscis and inflicted a wound, causing a sensation
similar to that produced by the burning of phosphorus under the skin.
The wound was a small, deep, triangular mark, succeeded by a
watery vesicle. The natives of New Guinea have a wholesome dread
of the bite of Cones. Mr. C. Hedley relates[149] that while collecting
on a coral reef he once rolled over a boulder and exposed a living C.
textile. Before he could pick it up, one of the natives hastily snatched
it away, and explained, with vivid gesticulations, its hurtful qualities.
On no account would he permit Mr. Hedley to touch it, but insisted on
himself placing it in the bottle of spirits.
Fig. 27.—A tooth from the

radula of Conus
imperialis L., × 50,
showing barb and
poison duct.
Mimicry and Protective Coloration.
Cases of Mimicry, or protective resemblance, when a species
otherwise defenceless adopts the outward appearance of a better
protected species, are rare among the Mollusca. Karl Semper[150]
mentions an interesting case of the mimicry of Helicarion tigrinus by
Xesta Cumingii, in the Philippines. It appears that all species of
Helicarion possess the singular property of shaking off the ‘tail’ or
hinder part of the foot, when seized or irritated. Specimens captured
by collectors, Hel. tigrinus amongst them, have succeeded in
escaping from the hand, and concealing themselves, by a sort of
convulsive leap, among the dry leaves on the ground. This power of
self-amputation must be of great value to Helicarion, not only as
enabling it to escape from the clutch of its enemies, but also as
tending to discourage them from attempting to capture it at all. Now
the genus Xesta is, in anatomy, very far removed from Helicarion,
and the majority of the species are also, as far as the shell is
concerned, equally distinct. Xesta Cumingii, however, has, according
to Semper, assumed the appearance of a Helicarion, the thin shell,
the long tail, and the mantle lobes reflected over the shell; but it has
not the power of parting with its tail at short notice. It lives associated
with Helicarion, and so close is the resemblance between them that,
until Semper pointed out its true position, it had always been
classified as a member of that group.
In the same passage Semper draws attention to two other cases
of apparent mimicry. The first is another species of Xesta
(mindanaensis) which closely resembles a species of Rhysota
(Antonii), a genus not indeed so far removed from Xesta as
Helicarion, but, as far as the shell is concerned, well distinguished
from it. In this case, however, there is no obvious advantage gained
by the resemblance, since Rhysota as compared with Xesta is not
known to possess any definite point of superiority which it would be
worth while to counterfeit. A second case of resemblance between
certain species of the genus Chloraea and the characteristic
Philippine group Cochlostyla will not hold good as affording evidence
of mimicry, for Chloraea is now recognised as a sub-genus of
Cochlostyla.
The Mollusca are not much mimicked by creatures of different
organisation. This appears at first sight strange, since it might have
been thought that the strong defensive house of a snail was worth
imitating. Still it is probably not easy for creatures bilaterally
symmetrical to curl themselves up into an elevated spiral for any
length of time. One or two instances, however, may be mentioned.
The larva of a moth belonging to the Psychidae, and occurring in
France, Germany, the Tyrol, and Syria, coils itself up into a sinistral
spiral of three whorls, and is aptly named Psyche helix, a kindred
species from Italy being known as Ps. planorbis.
An insect larva (Cochlophora valvata) from E. Africa is said to
resemble a Valvata or young Cyclostoma. In this case the spiral is
indifferently dextral or sinistral, the ‘shell’ being formed of masticated
vegetable matter, united together by threads spun by the larva.
Certain larvae of the Phryganeidae (“Caddis-worms”) enclose
themselves in houses which more or less resemble a spiral shell,
and have in some cases actually been described as molluscan; such
species, some of which belong to Helicopsyche, have been noticed
in S. Europe, Ceylon, Further India, China, Tasmania, New Zealand,
Tennessee, Mexico, Central America, Venezuela, Brazil, and
Argentina, and all[151] possess a dextral ‘shell.’ In all these cases
‘mimicry’ is probably not so much to be thought of as the practical
advantages which accrue to the animal in question from the spiral
form, which gives it greater strength to resist external blows, and
enables it to occupy, during a very defenceless portion of its
existence, a very small amount of space.
The larva of some species of the Syrphidae (Diptera) fixes itself
on the under side of stones in the Tyrol, and closely resembles a
small slug. The naturalist Von Spix, in 1825, described to the
Bavarian Academy as a new genus of land Mollusca a somewhat
similar larval form found in decaying wood on the banks of a German
lake.[152] Simroth mentions[153] a curious case as occurring near
Grimma. The caterpillars of certain Microlepidoptera occur on slabs
of porphyry, associated with a species of Clausilia. Besides being of
the same colour as the Clausiliae, the caterpillars have actually
developed cross lines on the back, i.e. on the side turned away from
the rock, in imitation of the suture of the mollusc.
It has been suggested[154] that there is mimicry between Aeolis
papillosa (a common British nudibranch) and Sagartia troglodytes
(an Actinian), and also between another species of Sagartia and
Aeolidiella Alderi. The facts observed are not sufficient to warrant a
decided opinion, but it seems more probable that the Actinian mimics
the nudibranch than vice versâ, since Aeolis is known to be
unpalatable to fishes.
Fig. 28.—A, Strombus mauritianus

Lam., which mimics Conus in
shape. B, Conus janus
Hwass, Mauritius.
Certain species of Strombus (mauritianus L., luhuanus L.) show a
remarkable similarity in the shape of the shell to that of Conus, so
much so, that a tiro would be sure to mistake them, at first sight, for
Cones. In the case of S. luhuanus at least, this similarity is increased
by the possession of a remarkably stout brown epidermis. Now
Conus is a flesh-eating genus, armed with very powerful teeth which
are capable of inflicting even on man a poisonous and sometimes
fatal wound (see p. 66). Strombus, on the other hand, is probably
frugivorous, and is furnished with weak and inoffensive teeth. It is
possible that this resemblance is a case of ‘mimicry.’ It is quite
conceivable that powerful fishes which would swallow a Strombus
whole and not suffer for it, might acquire a distaste for a Cone, which
was capable of lacerating their insides after being swallowed. And
therefore the more like a Cone the Strombus became, the better
chance it would have of being passed over as an ineligible article of
food.
Protective coloration is not uncommon among the Mollusca.
Littorina obtusata is habitually found, on our own coasts, on Fucus
vesiculosus, the air-bladders of which it closely resembles in colour
and shape. Littorina pagodus, a large and showy species, resembles
so closely the spongy crumbling rocks of Timor, on which it lives, that
it can hardly be discerned a pace off. Helcion pellucidum, the
common British ‘blue limpet,’ lives, when young, almost exclusively
on the iridescent leaves of the great Laminariae, with the hues of
which its own conspicuous blue lines harmonise exactly. In mature
life, when the Helcion invariably transfers its place of abode to the
lower parts of the stalk and finally to the root of the Laminaria, which
are quite destitute of iridescence, these blue lines disappear or
become much less marked.
The specimens of Purpura lapillus which occur at Newquay in
Cornwall are banded with rings of colour, especially with black and
white, in a more varied and striking way than any other specimens
that have ever occurred to my notice. I am inclined to refer this
peculiarity to a tendency towards protective coloration, since the
rocks on which the Purpura occurs are often banded with veins of
white and colour, and variegated to a very marked extent.
Ovula varies the colour of its shell from yellow to red, to match the
colour of the Gorgonia on which it lives. The same is the case with
Pedicularia, which occurs on red and yellow coral.
Helix desertorum, by its gray-brown colour, harmonises well with
the prevailing tint of the desert sands, among which it finds a home.
Benson observes that the gaudy H. haemastoma, which lives on the
trunks of palm-trees in Ceylon, daubs its shell with its excrement.
Our own Buliminus obscurus, which lives principally on the trunks of
smooth-barked trees, daubs its shell with mud, and must often
escape the observation of its enemies by its striking resemblance to
the little knots on the bark, especially of beech trees, its favourite
haunt. Some species of Microphysa, from the West Indies, habitually
encrust their shells with dirt, and the same peculiarity in Vitrina has
already been mentioned. Ariophanta Dohertyi Aldr., a recent
discovery from Sumatra, is of a green colour, with a singularly
delicate epidermis; it is arboreal in its habits, and is almost invisible
amongst the foliage.[155] Many of our own slugs, according to
Scharff, are coloured protectively according to their surroundings. A
claret-coloured variety of Arion ater occurred to this observer only in
pine woods, where it harmonised with the general colouring of the
ground and the pine-needles, while young winter forms of the same
species choose for hiding-places the yellow fallen leaves, whose
colour they closely resemble. Limax marginatus (= arborum Bouch.)
haunts tree trunks, and may easily be mistaken for a piece of bark;
Amalia carinata lives on and under the ground, and in colour
resembles the mould; Arion intermedius feeds almost exclusively on
fungi, to which its colour, which is white, gray, or light yellow, tends to
approximate it closely; Geomalacus maculosus conceals itself by its
striking resemblance to the lichens which grow on the surface of
rocks, and actually presumes on this resemblance so much as to
expose itself, contrary to the usual custom of its congeners, to the
full light of the afternoon sun.[156]
Several views have been advanced with regard to the dorsal
papillae, or cerata, in the Nudibranchs. Professor W. A. Herdman,
who has examined a considerable number of our own British
species, in which these processes occur, is of opinion[157] that they
are of two quite distinct kinds. In the first place, they may contain
large offshoots, or diverticula, of the liver, and thus be directly
concerned in the work of digestion. This is the case with Aeolis and
Doto. In the second place, they may be simply lobes on the skin,
with no connexion with the liver, and no special function to perform.
This is the case with Tritonia, Ancula, and Dendronotus.
Professor Herdman is of opinion that although the cerata may in
all cases aid in respiration to a certain extent, yet that extent is so
small as to be left out of consideration altogether. He regards the
cerata in both the two classes mentioned above as “of primary
importance in giving to the animals, by their varied shapes and
colours, appearances which are in some cases protective, and in
others conspicuous and warning.”
Thus, for instance, Tritonia plebeia, which is fairly abundant at
Puffin and Hilbre Is., appears always to be found creeping on the
colonies of a particular polyp, Alcyonium digitatum, and nowhere
else. The specimens in each colony of the polyp differ noticeably
both in the matter of colour, and of size, and of varied degrees of
expansion. The Tritonia differs also, being marked in varied tints of
yellow, brown, blue, gray, black, and opaque white, in such a way as
to harmonise with the varied colours of the Alcyonium upon which it
lives. The cerata on the back of the Tritonia contribute to this general
resemblance. They are placed just at the right distance apart, and
are just the right size and colour, to resemble the crown of tentacles
on the half-expanded polyp.
Similarly, Doto coronata, which, when examined by itself, is a very
conspicuous animal, with showy, bright-coloured cerata, is found by
Professor Herdman to haunt no other situations but the under side of
stones and overhanging ledges of rock which are colonised by a
hydroid, known as Clava multicornis. The Doto is masked by the
tentacles and clusters of sporosacs on the zoophyte, with whose
colouring and size its own cerata singularly correspond. A similar
and even more deceptive correspondence with environment was
noticed in the case of the very conspicuous Dendronotus
arborescens.
In these cases, the colouring and general shape of the cerata are
protective, i.e. they match their surroundings in such a way as to
enable the animal, in all probability, to escape the observation of its
enemies. According to Professor Herdman, however, the brilliant and
showy coloration of the cerata of Aeolis is not protective but
‘warning.’ Aeolis does not hide itself away as if shunning
observation, like Doto, Tritonia, and Dendronotus; on the contrary, it
seems perfectly fearless and indifferent to being noticed. Its cerata
are provided with sting-cells, like those of Coelenterata, at their tips,
and its very conspicuousness is a warning to its enemies that they

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Spatial Planning in Ghana Origins

Contemporary Reforms and Practices

Ecosystem Services and Green Infrastructure:

Experimental and Expanded Animation: New Perspectives

New Atheism Critical Perspectives and Contemporary

Sustainable Water Management New Perspectives Design

Ecosystem Services for Spatial Planning Silvia Ronchi

Trends in Spatial Analysis and Modelling: Decision-

Materiality and Managerial Techniques : New

Spatial Planning in Service Delivery: Towards

Series Advisory Editors

More information about this series at http://www.springer.com/series/14773

Spatial Planning in Ghana

ISSN 2365-757X ISSN 2365-7588 (electronic)

Library of Congress Control Number: 2018957058

© Springer Nature Switzerland AG 2019

In 2012, I had the unique opportunity to be involved in a new wave of institutional

Dublin, Ireland Ransford A. Acheampong, Ph.D.

I would like to thank Stephen Asabere, Priscilla Ankomah-Hackman and

Part I Setting the Scene

3.3 Landmark (Spatial) Plan-Making Projects in the Era

Part II Modern-Day Planning Systems and Scales

5 The Inception of Regional Spatial Planning . . . . . . . . . . . . . . . . .. 83

6.5 Non-Compliance with Local Plans and Development

Part III Issues in Spatial Planning

9.4 Urbanization and Urban Growth Management . . . . . . . . . . . . . 192

11.4 Towards Integrated Spatial Development and Transport Policy

13.3.3 Housing Supply—The Role of Spatial Planning . . . . . . 297

Ransford A. Acheampong Ph.D., M.Phil. Cambridge, UK; B.Sc. KNUST, Ghana

AMA Accra Metropolitan Area/Assembly

KMA Kumasi Metropolitan Area/Assembly

TAZs Trafﬁc Analysis Zones

Fig. 2.1 Constituents of a typical spatial planning system . . . . . . . . . . . . 21

Fig. 3.11 Three-tier model of spatial development frameworks and

Table 3.1 Periodizing signiﬁcant developments in the history of spatial

Box 6.1 Functions of the District Spatial Planning Committee and

Keywords Spatial planning · Institutions · Legal systems · Socio-economic

© Springer Nature Switzerland AG 2019 3

1 See, e.g., Silva (2015) and Bhan et al. (2018).

Enemies of the Mollusca

Fig. 24.—Illustrating the elaborate arrangement of teeth in the

Fig. 25.—Xenophora (Phorus)

Fig. 27.—A tooth from the

Fig. 28.—A, Strombus mauritianus

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