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ECOSYSTEM SERVICES FOR
WELL-BEING IN DELTAS
Integrated Assessment
for Policy Analysis
Edited by Robert J. Nicholls, Craig W. Hutton, W. Neil Adger,
Susan E. Hanson, Md. Munsur Rahman and Mashfiqus Salehin
Ecosystem Services for Well-Being in Deltas
 Robert J. Nicholls
Craig W. Hutton
W. Neil Adger • Susan E. Hanson
Md. Munsur Rahman
Mashfiqus Salehin
Editors

Ecosystem Services
for Well-Being
in Deltas
Integrated Assessment for Policy
Analysis
Editors
Robert J. Nicholls Craig W. Hutton
Faculty of Engineering and the Environment Geodata Institute
and Tyndall Centre for Climate Change Geography and Environment
Research, University of Southampton University of Southampton
Southampton, UK Southampton, UK

W. Neil Adger Susan E. Hanson

Geography, College of Life and Faculty of Engineering and the
Environmental Sciences Environment and Tyndall Centre for
University of Exeter Climate Change Research
Exeter, UK University of Southampton
Southampton, UK
Md. Munsur Rahman
Institute of Water and Flood Management Mashfiqus Salehin
Bangladesh University of Engineering and Institute of Water and Flood Management
Technology Bangladesh University of Engineering and
Dhaka, Bangladesh Technology
Dhaka, Bangladesh

ISBN 978-3-319-71092-1    ISBN 978-3-319-71093-8 (eBook)

https://doi.org/10.1007/978-3-319-71093-8

Library of Congress Control Number: 2018941812

© The Editor(s) (if applicable) and The Author(s) 2018. This book is an open access publication.
Open Access This book is licensed under the terms of the Creative Commons Attribution 4.0 International
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Printed on acid-free paper

This Palgrave Macmillan imprint is published by the registered company Springer International
Publishing AG part of Springer Nature.
The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
 Dedication: To Dr. Nazmul Haq, University of Southampton, aka ‘Uncle’,
for his facilitation of the research herein and his ongoing commitment to the
people of Bangladesh.
 Foreword

Deltas, Ecosystem Services and the Sustainable

Well-Being of Humans and the Rest of Nature
Humanity is finally rediscovering an important relationship—the inter-
dependent relationship between humans and the rest of nature. The
industrial revolution and some religious traditions have emphasised the
distinctions between humans and ‘nature’—that humans are somehow
above, apart from or fundamentally different from the rest of nature. In
fact, the more we learn about the way the world and its complex inter-
connected systems function, the more we recognise that homo sapiens is,
and has always been, an integral component of the ecosystems it is
embedded within. Humans are not apart from nature, but are a part of
the natural world, and their health and well-being cannot be understood
or managed separate from that complex and evolving context.
The concept of ecosystem services makes this interdependence with the
rest of nature more apparent and quantitative. It does this by analysing,
modelling, quantifying and valuing the degree to which humans are
­connected with and benefit from the ecosystems that enclose them.
Ecosystems provide a range of services that are of fundamental impor-
tance to human well-being, health, livelihoods and survival (Costanza
et al. 1997; Daily 1997; MEA 2005; de Groot et al. 2014).

vii
viii Foreword

The idea that preserving the environment as an asset, rather than an

impediment to economic and social development, is both very old and
very new. For most of human history, at least until the start of the Industrial
Revolution, the benefits humans derived from the rest of nature were well
recognised and embedded in various cultural rules and norms. Parts of
forests, lakes, wetlands or mountains were often deemed sacred and off
limits. But it is no coincidence that these sacred natural assets also supplied
essential life-support services for the communities involved. This is in stark
contrast to the post-industrial view in much of the Western world that
nature is merely a pretty picture—nice to enjoy if you can afford it but not
essential to the more important business of ‘growing the economy’. Too
often, when the issue of conservation of the environment has entered pub-
lic or political discussions, it has been purported to come at a cost, and the
discussion has been framed as ‘the environment versus the economy’.
Probably the most important contribution of the widespread recogni-
tion of ecosystem services is that it reframes the relationship between
humans and the rest of nature to be more consistent with what we know.
A better understanding of the role of ecosystem services emphasises our
natural assets as critical ingredients to inclusive wealth, well-being and
sustainability. Sustaining and enhancing human well-being requires a
balance of all of our assets—individual people, society, the built economy
and ecosystems. This reframing of the way we look at ‘nature’ is essential
to solving the problem of how to build a sustainable and desirable future
for humanity—a goal that we all share.
The ecosystem services concept makes it abundantly clear that the
choice of ‘the environment versus the economy’ is a false choice. If the
environment contributes significantly to human well-being, then it is a
major contributor to the real economy and the choice becomes how to
manage all our assets, including natural and human-made capital, more
effectively and sustainably (Costanza et al. 2000).
Interest in ecosystem services in both the research and policy commu-
nities has grown rapidly (Braat and de Groot 2012). As of this writing,
over 18,000 journal articles have been published on this topic, according
to SCOPUS, and the number is growing exponentially. The most highly
cited of these (with over 7,000 citations in SCOPUS as of this writing) is
one that I and 12 co-authors published in Nature in 1997 that estimated
 Foreword
   ix

the value of global ecosystem services to be in excess of US$33 trillion per

year, a figure larger than global gross domestic product (GDP) at the time
(Costanza et al. 1997). This admittedly crude underestimate, and a few
other early studies, stimulated a huge surge in interest in this topic. In
2005, the concept of ecosystem services gained broader attention when the
United Nations published its Millennium Ecosystem Assessment (MEA
2005). The MEA was a four-year, 1,300-scientist study for policymakers.
In 2008, a second international initiative was undertaken by the UN
Environment Programme, called The Economics of Ecosystems and
Biodiversity (TEEB 2010). The TEEB report was picked up extensively by
the mass media, bringing ecosystem services to a broader audience.
Hundreds of projects and groups are currently working towards better
understanding, modelling, valuation and management of ecosystem ser-
vices and natural capital. In 2012 the Intergovernmental Science-Policy
Platform on Biodiversity and Ecosystem Services (IPBES) was established.
IPBES is an intergovernmental body (similar to the IPCC) which provides
information on the state of biodiversity and ecosystem services for deci-
sion-making purposes. Its current membership includes 126 national gov-
ernments. Emerging global, national and regional networks like the
Ecosystem Services Partnership (www.es-partnership.org) have also
emerged. Ecosystem services are now poised to provide real solutions to the
problem of how to sustainably manage our critical natural capital assets.
From the perspective of ecosystem services, wetlands are among the
most important and valuable ecosystems in the world (de Groot et al.
2012). The recognition of this value is a far cry from the situation not
that long ago (and still prevalent in some places) when wetlands were
considered to be ‘wastelands’ and every effort was made to drain, fill and
convert them to other land uses.
Coastal wetlands, and in particular large river deltas, are especially
important and valuable. River deltas contain the majority of coastal wet-
lands. However, they are also among the most impacted by human activi-
ties and 1 in 14 people globally live in deltaic regions (Day et al. 2016).
The world’s most populated delta is the Ganges–Brahmaputra–Meghna
in Bangladesh.
This book is a compendium of some of the latest work on understand-
ing, valuing and managing ecosystem services in this, one of the most
x Foreword

important and vulnerable delta ecosystems in the world. It takes a much

needed ‘whole systems’ approach to understanding the current status and
trends in this complex system and focuses on the fundamental relation-
ships between the biophysical system and the welfare of the diverse
human communities that rely on it.
If we are to build the sustainable and desirable future we all want, we
need to be able to understand, model and value complex social-ecological
systems in the comprehensive way this book exemplifies. It is truly a
model that needs to be broadly emulated.
Robert Costanza
Professor and VC’s Chair in Public Policy
Crawford School of Public Policy
The Australian National University

References
Braat, L., and R. de Groot. 2012. The ecosystem services agenda: Bridging the
worlds of natural science and economics, conservation and development, and
public and private policy. Ecosystem Services 1: 4–15.
Costanza, R., R. d’Arge, R. de Groot, S. Farber, M. Grasso, B. Hannon,
K. Limburg, S. Naeem, R.V. Oneill, J. Paruelo, R. G. Raskin, P. Sutton, and
M. van den Belt. 1997. The value of the world’s ecosystem services and natu-
ral capital. Nature 387 (6630): 253–260.
Costanza, R., M. Daly, C. Folke, P. Hawken, C. S. Holling, A.J. McMichael,
D. Pimentel, and D. Rapport. 2000. Managing our environmental portfolio.
Bioscience 50 (2): 149–155.
Daily, G.C. (1997). Nature’s Services: Societal dependence on natural ecosys-
tems. Washington, DC: Island Press.
Day, J.W., J. Agboola, Z. Chen, C. D’Elia, D.L. Forbes, L. Giosan, P. Kemp,
C. Kuenzer, R.R. Lane, R. Ramachandran, J. Syvitski, and A. Yañez-
Arancibia. 2016. Approaches to defining deltaic sustainability in the 21st
century. Estuarine, Coastal and Shelf Science 183: 275–291.
de Groot, R., L. Brander, S, van der Ploeg, R. Costanza, F. Bernard, L. Braat,
M. Christie, N. Crossman, A. Ghermandi, L. Hein, S. Hussain, P. Kumar,
A. McVittie, R. Portela, L.C. Rodriguez, P. ten Brink, and P. van Beukering.
2012. Global estimates of the value of ecosystems and their services in mon-
etary units. Ecosystem Services 1: 50–61.
 Foreword
   xi

Millennium Ecosystem Assessment (MEA). 2005. Ecosystems and human well-

being: Synthesis. Washington DC: Island Press.
TEEB. 2010. Mainstreaming the economics of nature: A synthesis of the approach,
conclusions and recommendations of TEEB. London/Washington: Earthscan.
 Participants from the Following
Organisations Attended Workshops
Within the Project

Ministries of the Government of the People’s Republic of

Bangladesh

Department of Agricultural Extension (DAE)

Department of Fisheries (DoF)
Different Wings/Divisions of Planning Commission
General Economics Division of Planning Commission
Ministry of Environment and Forest (MoEF)
Ministry of Finance (MoF)
Ministry of Water Resources (MoWR)

Bangladesh Organisations

Bangladesh Agricultural Development Corporation (BADC)

Bangladesh Agricultural Research Institute (BARI)
Bangladesh Bureau of Statistics (BBS)
Bangladesh Institute of Development Studies (BIDS)
Bangladesh Rice Research Institute (BRRI)
Bangladesh Meteorological Department (BMD)
Bangladesh Water Development Board (BWDB)
Comprehensive Disaster Management Program (CDMP)

xiii
xiv Participants from the Following Organisations Attended…

Dhaka University (DU)

Institute of Water Modeling (IWM)
River Research Institute (RRI)
Soil Resources Development Institute (SRDI)
Space Research and Remote Sensing Organization (SPARRSO)
Water Resources Planning Organisation (WARPO)
WildTeam

Local Organisations

Cyclone Preparedness Programme

Local community-based organisations
Local Disaster Risk Reduction (DDR) Volunteers
Local Government Engineering Department (LGED)
Local media
Local mosque committee
Local non-governmental organisations
Local schools
Shelter Management Committee
Small entrepreneurship
Switch gate committee
Union Parishad
Upazila Administration
Upazila Parishad
Water Management Committee

International Organisations

Asian Development Bank

CARE
Delta Plan Consultants (Deltares, Bandudeltas)
Food and Agriculture Organization of the United Nations (FAO)
German Development Cooperation (GIZ)
Global Water Partnership
 Participants from the Following Organisations Attended…
   xv

International Organization for Migration (IOM)

International Union for Conservation of Nature (IUCN)
United Nations Development Programme (UNDP)
World Bank
World Food Program (WFP)
World Health Organization (WHO)
 Preface

Deltas provide diverse ecosystem services and benefits for their large pop-
ulations. At the same time, deltas are also recognised as one of the most
vulnerable coastal environments, with a range of drivers operating at mul-
tiple scales, from global climate change and sea-level rise to delta-­scale
subsidence and land cover change. Lastly, many delta populations experi-
ence significant poverty. Hence when the Ecosystem Services for Poverty
Alleviation (ESPA) programme was announced, we rapidly focussed on
deltas as an issue for study. The focus of the book is the world’s most
populated delta, the Ganges–Brahmaputra–Meghna Delta, and more
particularly within coastal Bangladesh west of the Lower Meghna River.
In our first visit to Dhaka, Bangladesh, in 2010, we recognised the com-
plexity and challenges of understanding rural livelihoods in a dynamic delta.
We held an intensive multidisciplinary workshop of UK and Bangladeshi
scientists, followed by an inspiring visit to the Sundarbans. The resulting
debates and conclusions, supported by acres of white board conceptual
maps, formed the foundation that became the ESPA Deltas (Assessing
Health, Livelihoods, Ecosystem Services and Poverty Alleviation in Populous
Deltas) international consortium project. This involves more than 120 indi-
viduals and 21 institutions across Bangladesh, India and the UK. The col-
lective thinking and experience of this team is distilled into this book, which
examines the present and future of ecosystem services and livelihoods in

xvii
xviii Preface

coastal Bangladesh. It reflects the strong commitment to integration and a

transdisciplinary approach, embracing disciplines as diverse as physical
oceanography, sediment dynamics, agriculture, demographics and poverty.
Input of policy experts and a substantial array of stakeholders are also fun-
damental. This study provided opportunities for substantial learning across
standard discipline boundaries, providing co-produced policy relevant out-
puts and insights. It also fostered a family of researchers who developed a
shared understanding that could be applied to this difficult and challenging
problem. This included effective sharing of knowledge and learning to ques-
tion and contribute effectively outside an individual’s specialist field.
Integration is core to what has been accomplished here bringing
together natural and social sciences in ways that are distinct and ground-­
breaking. Such integration needs to start as the research is initiated and is
an ongoing process. Integration needs to be core to the project with key
questions and themes that are properly resourced. To be policy relevant,
the research must be guided by the perspectives, needs and expertise
encapsulated by local stakeholders, especially the decision-making pro-
cesses and governance context of the deltas in question. Stakeholders
from civil society, the non-government sector and of course agencies of
government are all involved in policy formulation. Indeed, one of the
outstanding successes of the ESPA Deltas collaboration, which is reflected
in this book, has been the engagement with and the impact on the policy
context of Bangladesh. The research has raised, for the first time, consid-
eration of ecosystem services, their links to poverty and livelihoods and
their influence in the national policy and planning process across a range
of government agencies including the Government of Bangladesh,
Planning Commission and other government partners such as the Water
Resources Planning Organization (WARPO). Indeed the Government of
Bangladesh has requested continued engagement and further develop-
ment of some of the modelling tools in the context of the Bangladesh
Delta Plan 2100, which is a new national planning approach. Engaging
with policy was always a main aspiration of the research and is perhaps
the aspect of which we are most proud.
The research provides both the foundation and analyses which has led
to some of its most innovative approaches and significant insights. This
book offers an overarching and integrated framework to analyse changing
 Preface
   xix

ecosystem services in deltas and the implications for human well-being,

focussing in particular on the provisioning ecosystem services of agricul-
ture, inland and offshore capture fisheries, aquaculture and mangroves that
directly support livelihoods. Each chapter contributes to the wider inte-
grated assessment. Indeed, throughout the book there are reflections on
the process of integrating information on the different environmental,
social and economic dimensions of coastal management. The more detailed
work supports significant conclusions that challenge elements of the per-
ceived wisdom concerning human–environment relations and progress for
the future, under the Sustainable Development Goals. We highlight, for
example, that while ecosystem services support all populations in deltas,
they act as a more critical safety net for the poorest and most marginalised
delta populations. We show that while climate change has a real and tan-
gible impact on the coastal zone, demographic, social dynamics and policy
changes are likely to be more significant until at least 2050.
This book is not intended as a tool kit or specific guide to conducting
integrated research in deltas or major coastal systems throughout the
world. It offers, rather, a detailed account of a major integrative assess-
ment relevant to development dilemmas in major ecosystems where bio-
physical, ecological and social dimensions are strongly coupled. This
approach can be generalised beyond tropical deltas and even coastal
zones; it addresses fundamental questions regarding the relationship of
ecosystem services to the welfare of diverse rural communities that are
important in every corner of the world today.

University of Southampton Robert J. Nicholls

Southampton, UK  Craig W. Hutton
Susan E. Hanson
University of Exeter W. Neil Adger
Exeter, UK 
Bangladesh University of Engineering  Md. Munsur Rahman
and Technology Mashfiqus Salehin
Dhaka, Bangladesh 
 Acknowledgements

The book is a reflection on a diverse series of activities across biophysical,

ecological and social science research on delta systems. The research is the
outcome, firstly, of a major consortium project funded in the UK from
2012 to 2016. The research ‘ESPA Deltas (Assessing Health, Livelihoods,
Ecosystem Services and Poverty Alleviation in Populous Deltas)’
NE-J002755-1 was funded with support from the Ecosystem Services for
Poverty Alleviation (ESPA) programme. The ESPA programme was funded
by the Department for International Development (DFID), the Economic
and Social Research Council (ESRC) and the Natural Environment Research
Council (NERC). We thank them for this funding and their support,
including making this book an open access publication.
The ESPA Delta partnership involves researchers in Bangladesh, India and
the UK and across a wide range of relevant disciplines. In the UK these are
Universities of Southampton, Exeter, Dundee, Oxford, Bath, Plymouth
Marine Laboratory, the National Oceanography Centre Liverpool and the
Hadley Centre of the UK Met Office. The Bangladeshi partners are the
Institute of Water and Flood Management at the Bangladesh University of
Engineering and Technology, Bangladesh Agricultural Research Institute,
Technological Assistance for Rural Advancement, Ashroy Foundation,
Bangladesh Agricultural University, Bangladesh Institute of Development
Studies, Center for Environmental and Geographical Information Services,
Institute of Livelihood Studies, ­International Union for Conservation of
xxi
xxii Acknowledgements

Nature, Water Resources Planning Organization and International

Centre for Diarrhoeal Disease Research, Bangladesh. The project’s Indian
partners are the University of Jadavpur and the Indian Institute of
Technology Kanpur.
A project with this large scope and duration, and strong stakeholder
engagement, depends on the support of many individuals and organisa-
tions. They are too numerous to list comprehensively, but we acknowl-
edge their critical contribution, such as participating in the numerous
workshops and survey work that we organised.
In particular, the project team would also like to extend their thanks to
the following individuals and their organisations who provided essential
support during the research and greatly facilitated our engagement with
the national policy process:

• Professor Shamsul Alam, Member (Senior Secretary) General

Economics Division, Planning Commission, Government of the
People’s Republic of Bangladesh
• Dr. Sultan Ahmed, Director, Natural Resources Management,
Department of Environment (DoE), Government of the People’s
Republic of Bangladesh
• Dr. Taibur Rahman, Senior Assistant Chief, General Economics
Division, Planning Commission, Government of the People’s Republic
of Bangladesh
• Mr. Md. Mafidul Islam, Joint Chief, General Economics Division,
Planning Commission, Government of the People’s Republic of
Bangladesh

Last but not least, we also express our gratitude to the people of
Bangladesh who, during our research, provided hospitality and were ever
willing to give their time and enthusiasm to ensure that our research
remained relevant and successfully achieved its aim.
 Contents

Part 1 Research Highlights and Framework    1

1 Ecosystem Services, Well-Being and Deltas: Current

Knowledge and Understanding   3
W. Neil Adger, Helen Adams, Susan Kay, Robert J. Nicholls,
Craig W. Hutton, Susan E. Hanson, Md. Munsur Rahman,
and Mashfiqus Salehin

2 Ecosystem Services Linked to Livelihoods and Well-Being

in the Ganges-Brahmaputra-Meghna Delta  29
Helen Adams, W. Neil Adger, and Robert J. Nicholls

3 An Integrated Approach Providing Scientific

and Policy-­Relevant Insights for South-West Bangladesh  49
Robert J. Nicholls, Craig W. Hutton, Attila N. Lázár,
W. Neil Adger, Andrew Allan, Paul G. Whitehead, Judith Wolf,
Md. Munsur Rahman, Mashfiqus Salehin, Susan E. Hanson,
and Andres Payo

xxiii
xxiv Contents

4 Integrative Analysis for the Ganges-

Brahmaputra-­Meghna Delta, Bangladesh  71
Robert J. Nicholls, Craig W. Hutton, W. Neil Adger,
Susan E. Hanson, Md. Munsur Rahman,
and Mashfiqus Salehin

Part 2 Present Status of the Ganges-­Brahmaputra-­Meghna

Delta  91

5 Recent Trends in Ecosystem Services in Coastal

Bangladesh  93
John A. Dearing and Sarwar Hossain

6 Governance of Ecosystem Services Across Scales

in Bangladesh 115
Andrew Allan and Michelle Lim

7 Health, Livelihood and Well-Being in the

Coastal Delta of Bangladesh 131
Mohammed Mofizur Rahman and Sate Ahmad

8 Floods and the Ganges-Brahmaputra-­Meghna Delta 147

Anisul Haque and Robert J. Nicholls

Part 3 Scenarios for Policy Analysis 161

9 Integrating Science and Policy Through

Stakeholder-­Engaged Scenarios 163
Emily J. Barbour, Andrew Allan, Mashfiqus Salehin,
John Caesar, Robert J. Nicholls, and Craig W. Hutton

10 Incorporating Stakeholder Perspectives in Scenario

Development 179
Andrew Allan, Michelle Lim, and Emily J. Barbour
 Contents
   xxv

11 Regional Climate Change over South Asia 207

John Caesar and Tamara Janes

12 Future Scenarios of Economic Development 223

Alistair Hunt

Part 4 Observations and Potential Trends 247

13 Biophysical Modelling of the Ganges, Brahmaputra,

and Meghna Catchment 249
Paul G. Whitehead

14 Marine Dynamics and Productivity in the Bay of Bengal 263

Susan Kay, John Caesar, and Tamara Janes

15 A Sustainable Future Supply of Fluvial Sediment

for the Ganges-Brahmaputra Delta 277
Stephen E. Darby, Robert J. Nicholls, Md. Munsur Rahman,
Sally Brown, and Rezaul Karim

16 Present and Future Fluvial, Tidal and Storm Surge

Flooding in Coastal Bangladesh 293
Anisul Haque, Susan Kay, and Robert J. Nicholls

17 Modelling Tidal River Salinity in Coastal Bangladesh 315

Lucy Bricheno and Judith Wolf

18 Mechanisms and Drivers of Soil Salinity

in Coastal Bangladesh 333
Mashfiqus Salehin, Shahad Mahabub Chowdhury, Derek
Clarke, Shahjahan Mondal, Sara Nowreen, Mohammad
Jahiruddin, and Asadul Haque

19 Population Dynamics in the South-West of Bangladesh 349

Sylvia Szabo, Sate Ahmad, and W. Neil Adger
xxvi Contents

20 Land Cover and Land Use Analysis in Coastal Bangladesh  367

Anirban Mukhopadhyay, Duncan D. Hornby, Craig W. Hutton,
Attila N. Lázár, Fiifi Amoako Johnson, and Tuhin Ghosh

21 A Geospatial Analysis of the Social, Economic

and Environmental Dimensions and Drivers of Poverty
in South-West Coastal Bangladesh 383
Fiifi Amoako Johnson and Craig W. Hutton

22 Defining Social-Ecological Systems in South-West

Bangladesh 405
Helen Adams, W. Neil Adger, Munir Ahmed, Hamidul Huq,
Rezaur Rahman, and Mashfiqus Salehin

23 Characterising Associations between Poverty

and Ecosystem Services 425
Helen Adams, W. Neil Adger, Sate Ahmad, Ali Ahmed, Dilruba
Begum, Mark Chan, Attila N. Lázár, Zoe Matthews,
Mohammed Mofizur Rahman, and Peter Kim Streatfield

Part 5 Present and Future Ecosystem Services 445

24 Prospects for Agriculture Under Climate Change

and Soil Salinisation 447
Derek Clarke, Attila N. Lázár, Abul Fazal M. Saleh, and
Mohammad Jahiruddin

25 Marine Ecosystems and Fisheries: Trends and Prospects 469

Manuel Barange, Jose A. Fernandes, Susan Kay,
Mostafa A. R. Hossain, Munir Ahmed, and Valentina Lauria

26 Dynamics of the Sundarbans Mangroves in Bangladesh

Under Climate Change 489
Anirban Mukhopadhyay, Andres Payo, Abhra Chanda,
Tuhin Ghosh, Shahad Mahabub Chowdhury,
and Sugata Hazra
 Contents
   xxvii

27 Hypertension and Malnutrition as Health

Outcomes Related to Ecosystem Services 505
Ali Ahmed, Mahin Al Nahian, Craig W. Hutton,
and Attila N. Lázár

Part 6 Integration and Dissemination 523

28 Integrative Analysis Applying the Delta Dynamic

Integrated Emulator Model in South-West Coastal
Bangladesh 525
Attila N. Lázár, Andres Payo, Helen Adams, Ali Ahmed,
Andrew Allan, Abdur Razzaque Akanda, Fiifi Amoako Johnson,
Emily J. Barbour, Sujit Kumar Biswas, John Caesar, Alexander
Chapman, Derek Clarke, Jose A. Fernandes, Anisul Haque,
Mostafa A. R. Hossain, Alistair Hunt, Craig W. Hutton, Susan
Kay, Anirban Mukhopadhyay, Robert J. Nicholls, Abul Fazal
M. Saleh, Mashfiqus Salehin, Sylvia Szabo, and Paul G.
Whitehead

29 Communicating Integrated Analysis Research Findings 575

Mashrekur Rahman and Md. Munsur Rahman

Glossary of Bengali Words 587

Index 589
 Notes on Contributors

Helen Adams Department of Geography, King’s College London, London,

UK
Munir Ahmed TARA, Dhaka, Bangladesh
Mahin Al Nahian Climate Change and Health, International Center for
Diarrheal Disease Research, Bangladesh (icddr,b), Dhaka, Bangladesh
W. Neil Adger Geography, College of Life and Environmental Sciences,
University of Exeter, Exeter, UK
Sate Ahmad Faculty of Agricultural and Environmental Sciences, University of
Rostock, Rostock, Germany
Ali Ahmed Climate Change and Health, International Center for Diarrheal
Disease Research, Bangladesh (icddr,b), Dhaka, Bangladesh
Abdur Razzaque Akanda Irrigation and Water Management Institute,
Bangladesh Agricultural Research Institute, Gazipur, Bangladesh
Andrew Allan School of Law, University of Dundee, Dundee, UK
Manuel Barange Food and Agriculture Organization of the United Nations,
Rome, Italy
Plymouth Marine Laboratory, Plymouth, UK

xxix
xxx Notes on Contributors

Emily J. Barbour School of Geography and the Environment, University of

Oxford, Oxford, UK
Dilruba Begum Climate Change and Health, International Center for
Diarrheal Disease Research, Bangladesh (icddr,b), Dhaka, Bangladesh
Sujit Kumar Biswas Irrigation and Water Management Institute, Bangladesh
Agricultural Research Institute, Gazipur, Bangladesh
Lucy Bricheno National Oceanographic Centre, Liverpool, UK
Sally Brown Faculty of Engineering and the Environment and Tyndall Centre
for Climate Change Research, University of Southampton, Southampton, UK
John Caesar Met Office Hadley Centre for Climate Science and Services,
Exeter, Devon, UK
Mark Chan Department of Geography, King’s College London, London, UK
Abhra Chanda School of Oceanographic Studies, Jadavpur University, Kolkata,
India
Alexander Chapman Faculty of Engineering and the Environment and
Tyndall Centre for Climate Change Research, University of Southampton,
Southampton, UK
Shahad Mahabub Chowdhury International Union for Conservation of
Nature, Dhaka, Bangladesh
Derek Clarke Faculty of Engineering and the Environment and Tyndall Centre
for Climate Change Research, University of Southampton, Southampton, UK
Stephen E. Darby Geography and Environment, Faculty of Social, Human
and Mathematical Sciences, University of Southampton, Southampton, UK
John A. Dearing Geography and Environment, Faculty of Social, Human and
Mathematical Sciences, University of Southampton, Southampton, UK
Jose A. Fernandes Plymouth Marine Laboratory, Plymouth, UK
Tuhin Ghosh School of Oceanographic Studies, Jadavpur University, Kolkata,
India
Susan E. Hanson Faculty of Engineering and the Environment and
Tyndall Centre for Climate Change Research, University of Southampton,
Southampton, UK
 Notes on Contributors
   xxxi

Asadul Haque Department of Soil Science, Patuakhali Science and Technology

University, Patuakhali, Bangladesh
Anisul Haque Institute of Water and Flood Management, Bangladesh
University of Engineering and Technology, Dhaka, Bangladesh
Sugata Hazra School of Oceanographic Studies, Jadavpur University, Kolkata,
India
Duncan D. Hornby Geodata Institute, Geography and Environment,
University of Southampton, Southampton, UK
Mostafa A. R. Hossain Department of Fisheries Biology and Genetics,
Bangladesh Agricultural University, Mymensingh, Bangladesh
Sarwar Hossain Institute of Geography, University of Bern, Bern, Switzerland
Alistair Hunt Department of Economics, University of Bath, Bath, UK
Hamidul Huq Institute of Livelihood Studies, Bangladesh University of
Engineering and Technology, Dhaka, Bangladesh
Craig W. Hutton Geodata Institute, Geography and Environment, University
of Southampton, Southampton, UK
Mohammad Jahiruddin Department of Soil Science, Bangladesh Agricultural
University, Mymensingh, Bangladesh
Fiifi Amoako Johnson Social Statistics and Demography, Faculty of Social,
Human and Mathematical Sciences, University of Southampton, Southampton,
UK
Tamara Janes Met Office Hadley Centre for Climate Science and Services,
Exeter, Devon, UK
Rezaul Karim Institute of Water and Flood Management, Bangladesh
University of Engineering and Technology, Dhaka, Bangladesh
Susan Kay Plymouth Marine Laboratory, Plymouth, UK
Valentina Lauria Plymouth Marine Laboratory, Plymouth, UK
Attila N. Lázár Faculty of Engineering and the Environment and Tyndall
Centre for Climate Change Research, University of Southampton, Southampton,
UK
Michelle Lim Adelaide Law School, University of Adelaide, Adelaide, SA,
Australia
xxxii Notes on Contributors

Zoe Matthews Social Statistics and Demography, Faculty of Social, Human

and Mathematical Sciences, University of Southampton, Southampton, UK
Shahjahan Mondal Institute of Water and Flood Management, Bangladesh
University of Engineering and Technology, Dhaka, Bangladesh
Anirban Mukhopadhyay School of Oceanographic Studies, Jadavpur
University, Kolkata, India
Robert J. Nicholls Faculty of Engineering and the Environment and
Tyndall Centre for Climate Change Research, University of Southampton,
Southampton, UK
Sara Nowreen Institute of Water and Flood Management, Bangladesh
University of Engineering and Technology, Dhaka, Bangladesh
Andres Payo British Geological Survey, Keyworth, Nottingham, UK
Mashrekur Rahman Institute of Water and Flood Management, Bangladesh
University of Engineering and Technology, Dhaka, Bangladesh
Md. Munsur Rahman Institute of Water and Flood Management, Bangladesh
University of Engineering and Technology, Dhaka, Bangladesh
Mohammed Mofizur Rahman International Center for Diarrheal Disease
Research, Bangladesh (icddr,b), Dhaka, Bangladesh
Rezaur Rahman Institute of Water and Flood Management, Bangladesh
University of Engineering and Technology, Dhaka, Bangladesh
Abul Fazal M. Saleh Institute of Water and Flood Management, Bangladesh
University of Engineering and Technology, Dhaka, Bangladesh
Mashfiqus Salehin Institute of Water and Flood Management, Bangladesh
University of Engineering and Technology, Dhaka, Bangladesh
Peter Kim Streatfield Formerly of the International Center for Diarrheal
Disease Research, Bangladesh (icddr,b), Dhaka, Bangladesh
Sylvia Szabo Department of Development and Sustainability, Asian Institute
of Technology, Bangkok, Thailand
Paul G. Whitehead School of Geography and the Environment, University of
Oxford, Oxford, UK
Judith Wolf National Oceanographic Centre, Liverpool, UK
 List of Figures

Fig. 1.1 Interventions and processes related to ecosystem service

provision in delta environments 12
Fig. 1.2 Principal ecosystem processes and services in delta environ-
ments13
Fig. 1.3 Social-ecological systems and decision-making over
ecosystem services 15
Fig. 1.4 Guide to the individual and integrated research discussed in
this book 19
Fig. 2.1 Stylised representations of selected movements of people
and trade-offs between ecosystem services, between
social-ecological systems in the coastal zone of Bangladesh,
geographically and between seasons. The y-axis represents
the degree to which the services of that social-ecological
system are privately owned. The x-axis represents whether
the productivity of the social-ecological system is depen-
dent on the degradation of another. Arrows show flows of
labour, materials and process linkages. The diagrams
illustrate the inter-dependent nature of social-ecological
systems in the delta and the potential trade-offs in produc-
tivity and rural employability 40
Fig. 3.1 Conceptual view of the integrated model, including the
roles of stakeholders, domain experts, specialist integrators
within the research team and operators 52

xxxiii
xxxiv List of Figures

Fig. 4.1 Catchment area of the Ganges, Brahmaputra and Meghna

rivers (left), and major distributaries, seven national Divisions
(of Bangladesh) and urban centres (red dots) (right) 73
Fig. 4.2 The study area is located within the Coastal Zone of
Bangladesh (left) and is comprised of nine districts (right) 74
Fig. 4.3 Scales of analysis in this book 82
Fig. 4.4 The overall approach and flow of information in the analysis 83
Fig. 4.5 An iterative learning loop using ΔDIEM for policy analysis,
comprising (1) scenario development, including adaptation
responses, (2) qualitative to quantitative translation to
ΔDIEM inputs, (3) simulations using ΔDIEM and (4)
stakeholder review of the simulations, which can lead to a
new cycle of analysis (Reprinted with permission from
Nicholls et al. 2016) 84
Fig. 5.1 Provisioning, regulating and habitat ecosystem services over
recent decades: (a) total rice production, (b) total crop
production, (c) total fish production, (d) shrimp production,
(e) aquacultural production from natural wetlands and
artificial ponds, (f) production of natural forest materials, (g)
monitored salinity in the Poshur river at Mongla (Khulna),
(h) mean annual river discharge in the Lower Meghna-
Ganges river at Hardinge Bridge, (i) monitored depth to
groundwater at Dacope (Khulna), (j) average monitored soil
salinity at seven sites in Khulna and Patuakhali, (k) moni-
tored tree numbers at an experimental plot in the
Sundarbans, (l) estimated volume of growing mangrove in
the Sundarbans (see Hossain et al. 2016a for data sources) 96
Fig. 5.2 Selected indicators of environmental drivers: (a) mean
annual temperature, (b) mean annual rainfall, (c) averaged
annual sea level curve (Hiron Point and Khepupara;
Brammer 2014), (d) damage to crops, (e) cyclone frequency
and intensity (see Hossain et al. 2016a for data sources) 99
Fig. 5.3 Selected indicators of well-being: (a) total regional GDP,
(b) share of GDP across land production sectors, (c) lower
and higher poverty level, (d) infant and child mortality, (e)
trained medical care at births, (f) improved and unim-
proved sanitation, (g) improved and unimproved drinking
water, (h) uptake of education (see Hossain et al. 2016a for
data sources) 104
Another random document with
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 Fig. 20.—Various forms of Heliozoa. In 3, a is the entire animal and b the
flagellula; c.vac, contractile vacuole; g, gelatinous investment; nu, nucleus;
psd, pseudopodia; sk, siliceous skeleton; sp, spicules. (From Parker and
Haswell, after other authors.)

Each of these divides, and the two sister cells then conjugate after
the same fashion as in Actinophrys, but the nuclear divisions to form
the coupling nucleus are two in number, i.e. the nucleus divides into
two, one of which goes to the surface as the first polar body, and the
sister of this again divides to form a second polar body (which also
passes to the surface) and a pairing nucleus.[84] The two cells then
fuse completely, and surround themselves with a second gelatinous
cyst wall, separated from the outer one by a layer of siliceous
spicules. The nucleus appears to divide at least twice before the
young creep out, to divide immediately into as many Actinophrys-like
cells as there were nuclei; then each of these multiplies its nuclei, to
become apocytial like the adult form.
 Fig. 21.—Diagram illustrating the conjugation of Actinosphaerium. 1, Original
cell; 2, nucleus divides to form two, N2N2; 3, each nucleus again divides to
form two, N3 and n3, the latter passing out with a little cytoplasm as an
abortive cell; 4, repetition of the same process as in 3; 5, the two nuclei N4
have fused in syngamy to form the zygote nucleus Nz.

Schaudinn admits 24 genera (and 7 doubtful) and 41 species (and

18 doubtful). None are known fossil. Their geographical distribution
is cosmopolitan, as is the case with most of the minute fresh-water
Protista; 8 genera are exclusively marine, and Orbulinella has only
been found in a salt-pond; Actinophrys sol is both fresh-water and
marine, and Actinolophus has 1 species fresh-water, the other
marine. One of the 14 species of Acanthocystis is marine; the
remaining genera and species are all inhabitants of fresh water.[85]

4. Radiolaria
Sarcodina with the protoplasm divided by a perforated chitinous
central capsule into a central mass surrounding the nucleus, and an
outer layer; the pseudopodia radiate, never anastomosing enough to
form a marked network; skeleton either siliceous, of spicules, or
perforated; or of definitely arranged spicules of proteid matter
(acanthin), sometimes also coalescing into a latticed shell;
reproduction by fission and by zoospores formed in the central
capsule. Habitat marine, suspended at the surface (plankton), at
varying depths (zonarial), or near the bottom (abyssal).

Fig. 22.—Collozoum inerme. A, B, C, three forms of colony; D, small colony with

central capsules (c.caps), containing nuclei, and alveoli (vac) in ectoplasm;
 E, isospores, with crystals (c); F, anisospores; nu, nucleus. (From Parker
and Haswell.)

The following is Haeckel's classification of the Radiolaria:—

I. Porulosa (Holotrypasta).—Homaxonic, or nearly so. Central capsule
spherical in the first instance; pores numerous, minute, scattered; mostly
pelagic.
A. Spumellaria (Peripylaea).—Pores evenly scattered; skeleton of solid
siliceous spicules, or continuous, and reticulate or latticed, rarely absent;
nucleus dividing late, as an antecedent to reproduction.

B. Acantharia (Actipylaea).—Pores aggregated into distinct areas;

skeleton of usually 20 centrogenous, regularly radiating spines of acanthin,
whose branches may coalesce into a latticed shell; nucleus dividing early.

II. Osculosa (Monotrypasta).—Monaxonic; pores of central capsule limited

to the basal area (osculum), sometimes accompanied by two (or more)
smaller oscula at apical pole, mostly zonarial or abyssal.
C. Nassellaria (Monopylaea).—Central capsule ovoid, of a single layer;
pores numerous on the operculum or basal field; skeleton siliceous, usually
with a principal tripod or calthrop-shaped spicule passing, by branching,
into a complex ring or a latticed bell-shaped shell; nucleus eccentric, near
apical pole.

D. Phaeodaria (Cannopylaea, Haeck.; Tripylaea, Hertw.).—Central

capsule spheroidal, of two layers, in its outer layer an operculum, with
radiate ribs and a single aperture, beyond which protrudes the outer layer;
osculum basal, a dependent tube (proboscis); accessory oscula, when
present, simpler, usually two placed symmetrically about the apical pole;
skeleton siliceous, with a combination of organic matter, often of hollow
spicules; nucleus sphaeroidal, eccentric; extracapsular protoplasm
containing an accumulation of dusky pigment granules ("phaeodium").
 Fig. 23.—Actinomma asteracanthion. A, the shell with portions of the two outer
spheres broken away; B, section showing the relations of the skeleton to
the animal, cent.caps, Central capsule; ex.caps.pr, extra-capsular
protoplasm: nu, nucleus; sk.1, outer, sk.2, middle, sk.3, inner sphere of
skeleton. (From Parker and Haswell, after Haeckel and Hertwig.)

A. Spumellaria.

Sublegion (1). Collodaria.[86]—Skeleton absent or of detached spicules;

colonial or simple.
Order i. Colloidea.—Skeleton absent. (Families 1, 2.) Thalassicolla Huxl.;
Thalassophysa Haeck.; Collozoum Haeck.; Collosphaera J. Müll.; Actissa
Haeck.

Order ii. Beloidea.—Skeleton spicular. (Families 3, 4.)

Sublegion (2). Sphaerellaria.—Skeleton continuous, latticed or spongy,

reticulate.
Order iii. Sphaeroidea.—Skeleton of one or several concentric spherical
shells; sometimes colonial. (Families 5-10.) Haliomma Ehrb.; Actinomma
Haeck. (Fig. 23).

Order iv. Prunoidea.—Skeleton a prolate sphaeroid or cylinder, sometimes

constricted towards the middle, single or concentric. (Families 11-17.)

Order v. Discoidea.—Shell flattened, of circular plan, simple or concentric,

rarely spiral. (Families 18-23.)

Order vi. Larcoidea.—Shell ellipsoidal, with all three axes unequal or

irregular, sometimes becoming spiral. (Families 24-32.)[87]
 Fig. 24.—Xiphacantha (Acantharia). From the surface. The skeleton only, × 100,
(From Wyville Thomson.)

B. Acantharia.

Order vii. Actinelida.—Radial spines numerous, more than 20, usually

grouped irregularly. (Families 33-35.) Xiphacantha Haeck.

Order viii. Acanthonida.—Radial spines equal. (Families 36-38.)

Order ix. Sphaerophracta.—Radial spines 20, with a latticed spherical

shell, independent of, or formed from the reticulations of the spines.
(Families 39-41.) Dorataspis Haeck. (Fig. 25, A).

Order x. Prunophracta.—Radial spines 20, unequal; latticed shell,

ellipsoidal, lenticular, or doubly conical. (Families 42-44.)

C. Nassellaria.

Order xi. Nassoidea.—Skeleton absent. (Family 45.)

Order xii. Plectoidea.—Skeleton of a single branching spicule, the

branches sometimes reticulate, but never forming a latticed shell or a
sagittal ring. (Families 46-47.)

Order xiii. Stephoidea.—Skeleton with a sagittal ring continuous with the

branched spicule, and sometimes other rings or branches. (Families 48-
 51.) Lithocercus Théel (Fig. 26, A).

Order xiv. Spyroidea.—Skeleton with a latticed shell developed around the

sagittal ring (cephalis), and constricted in the sagittal plane, with a lower
chamber (thorax) sometimes added. (Families 52-55.)

Order xv. Botryoidea.—As in Spyroidea, but with the cephalis 3-4 lobed;
lower chambers, one or several successively formed. (Families 56-58.)

Order xvi. Cyrtoidea.—Shell as in the preceding orders, but without lobing

or constrictions. (Families 59-70.) Theoconus Haeck. (Fig. 25, B).

D. Phaeodaria.

Order xvii. Phaeocystina.—Skeleton 0 or of distinct spicules; capsule

centric. (Families 71-73.) Aulactinium Haeck. (Fig. 26, B).

Order xviii. Phaeosphaeria.—Skeleton a simple or latticed sphere, with no

oral opening (pylome); capsule central. (Families 74-77.)

Order xix. Phaeogromia.—Skeleton a simple latticed shell with a pylome

at one end of the principal axis; capsule excentric, sub-apical. (Families 78-
82.) Pharyngella Haeck.; Tuscarora Murr.; Haeckeliana Murr. (Fig. 28).

Order xx. Phaeoconchia.—Shell of two valves, opening in the plane

("frontal") of the three openings of the capsule. (Families 83-85.)

We exclude Haeckel's Dictyochida, with a skeleton recalling that of

the Stephoidea, but of the impure hollow substance of the
Phaeodaria (p. 84). They rank now as Silicoflagellates (p. 114).

The Radiolarian is distinguished from all other Protozoa by the

chitinous central capsule, so that its cytoplasm is separated into an
outer layer, the extracapsular protoplasm (ectoplasm), and a central
mass, the intracapsular, containing the nucleus.[88]
The extracapsular layer forms in its substance a gelatinous mass, of
variable reaction, through which the plasma itself ramifies as a
network of threads ("sarcodictyum"), uniting at the surface to
constitute the foundation for the pseudopodia. This gelatinous matter
constitutes the "calymma." It is largely vacuolated, the vacuoles
("alveoli"), of exceptional size, lying in the nodes of the plasmic
network, and containing a liquid probably of lower specific gravity
than seawater; and they are especially abundant towards the
surface, where they touch and become polygonal. On mechanical
irritation they disappear, to be formed anew after an interval, a fact
that may explain the sinking from the surface in disturbed water. This
layer may contain minute pigment granules, but the droplets of oil
and of albuminous matter frequent in the central layer are rare here.
The "yellow cells" of a symbiotic Flagellate or Alga, Zooxanthella, are
embedded in the jelly of all except Phaeodaria, and the whole
ectosarc has the average consistency of a firm jelly.

The pseudopodia are long and radiating, with a granular external

layer, whose streaming movements are continuous with those of the
inner network. In the Acantharia they contain a firm axial filament,
like that of the Heliozoa, which is traceable to the central capsule;
and occasionally a bundle of pseudopodia may coalesce to form a
stout process like a flagellum ("sarcoflagellum"). Here, too, each
spine, at its exit from the jelly, is surrounded by a little cone of
contractile filaments, the myophrisks, whose action seems to be to
pull up the jelly and increase the volume of the spherical body so as
to diminish its density.

Fig. 25.—Skeletons of Radiolaria. A, Dorataspis; B, Theoconus. (After Haeckel.)

The intracapsular protoplasm is free from Zooxanthella except in the
Acantharia. It is less abundantly vacuolated, and is finely granular. In
the Porulosa it shows a radial arrangement, with pyramidal stretches
of hyaline plasma separated by intervals rich in granules. Besides
the alveoli with watery contents, others are present with albuminoid
matter in solution. Oil-drops, often brilliantly coloured, occur either in
the plasma or floating in either kind of vacuole; and they are often
luminous at night. Added to these, the intracapsular plasm contains
pigment-granules, most frequently red or orange, passing into yellow
or brown, though violet, blue, and green also occur. The
"phaeodium,"[89] however, that gives its name to the Phaeodaria, is
an aggregate of dark grey, green, or brown granules which are
probably formed in the endoplasm, but accumulate in the
extracapsular plasm of the oral side of the central capsule. Inorganic
concretions and crystals are also found in the contents of the central
capsule, as well as aggregates of unknown composition, resembling
starch-grains in structure.

In the Monopylaea, or Nassellaria (Figs. 25, B, 26, A), the

endoplasm is differentiated above the perforated area of the central
capsule into a cone of radiating filaments termed the "porocone,"
which may be channels for the communication between the
exoplasm and the endoplasm, or perhaps serve, as Haeckel
suggests, to raise, by their contraction, the perforated area: he
compares them to the myophane striae of Infusoria. In the
Phaeodaria (Fig. 26, B), a radiating laminated cone is seen in the
outermost layer of the endoplasm above the principal opening
("astropyle"), and a fibrillar one around the two accessory ones
("parapyles"); and in some cases, continuous with these, the whole
outer layer of the endoplasm shows a meridional striation.

The nucleus is contained in the endoplasm, and is always at first

single, though it may divide again and again. The nuclear wall is a
firm membrane, sometimes finely porous. If there are concentric
shells it at first occupies the innermost, which it may actually come to
enclose, protruding lobes which grow through the several
perforations of the lattice-work, finally coalescing outside completely,
so as to show no signs of the joins. In the Nassellaria a similar
process usually results in the formation of a lobed nucleus,
contained in an equally lobed central capsule. The chromatin of the
nucleus may be concentrated into a central mass, or distributed into
several "nucleoli," or it may assume the form of a twisted, gut-like
filament, or, again, the nuclear plasm may be reticulated, with the
chromatin deposited at the nodes of the network.

Fig. 26.—A, Lithocercus annularis, with sagittal ring (from Parker and Haswell).
B, Aulactinium actinastrum. C, calymma; cent.caps., km, central capsule;
Ext.caps.pr., Extracapsular, and Int.caps.pr., intracapsular protoplasm; n,
nu, nucleus; op, operculum; ph, phaeodium; psd, pseudopodium; Skel.,
skeleton; z, Zooxanthella. (From Lang's Comparative Anatomy, after
Haeckel.)

The skeleton of this group varies, as shown in our conspectus, in the

several divisions.[90] The Acantharia (Figs. 24, 25, A) have a
skeleton of radiating spines meeting in the centre of figure of the
endoplasm, and forcing the nucleus to one side. The spines are
typically 20 in number, and emerge from the surface of the regular
spherical forms (from which the others may be readily derived)
radially, in five sets of four in the regions corresponding to the
equator and the tropics and polar circles of our world. The four rays
of adjacent circles alternate, so that the "polar" and "equatorial" rays
are on one set of meridians 90° apart, and the "tropical" spines are
on the intermediate meridians, as shown in the figures. By tangential
branching, and the meeting or coalescence of the branches,
reticulate (Figs. 23, 24, 25) and latticed shells are formed in some
families, with circles of openings or pylomes round the bases of the
spines. In the Sphaerocapsidae the spines are absent, but their
original sites are inferred from the 20 circles of pylomes.

In the Spumellaria the simplest form of the (siliceous) skeleton is that

of detached spicules, simple or complex, or passing into a latticed
shell, often with one or more larger openings (pylomes). Radiating
spines often traverse the whole of the cavity, becoming continuous
with its latticed wall, and bind firmly the successive zones when
present (Fig. 23).

Calcaromma calcarea was described by Wyville Thomson as having

a shell of apposed calcareous discs, and Myxobrachia, by Haeckel,
as having collections of the calcareous Coccoliths and
Coccospheres. In both cases we have to do with a Radiolarian not
possessing a skeleton, but retaining the undigested shells of its food,
in the former case (Actissa) in a continuous layer, in the latter
(Thalassicolla) in accumulations that, by their weight, droop and pull
out the lower hemisphere into distinct arms.

The (siliceous) skeleton of the Nassellaria is absent only in the

Nassoidea, and is never represented by distinct spicules. Its simplest
form is a "tripod" with the legs downward, and the central capsule
resting on its apex. The addition of a fourth limb converts the tripod
into a "calthrop," the central capsule in this case resting between the
upturned leg and two of the lower three regarded as the
"anterolateral"; the odd lower leg, like the upturned one, being
"posterior." Again, the skeleton may present a "sagittal ring," often
branched and spiny (Fig. 26, A), or combined with the tripod or
calthrop, or complicated by the addition of one or more horizontal
rings. Another type is presented by the "latticed chamber"
surrounding the central capsule, with a wide mouth ("pylome") below.
This is termed the "cephalis"; it may be combined in various ways
with the sagittal ring and the tripod or calthrop; and, again, it may be
prolonged by the addition of one, two, or three chambers below, the
last one opening by a pylome (Fig. 25, B). These are termed
"thorax," "abdomen," and "post-abdomen" respectively.

In the Phaeodaria the skeleton may be absent, spicular (of loose or

connected spicules) or latticed, continuous or bivalve. It is composed
of silica combined with organic matter, so that it chars when heated,
is more readily dissolved, and is not preserved in fossilisation. The
spicules or lattice-work are hollow, often with a central filament
running in the centre of the gelatinous contents. The latticed
structure of the shell of the Challengeridae (Fig. 28) is so fine as to
recall that of the Diatomaceae. In the Phaeoconchida the shell is in
two halves, parted along the "frontal" plane of the three apertures of
the capsule.

Fig. 27.—Scheme of various possible skeletal forms deposited in the meshes of

an alveolar system, most of which are realised in the Radiolaria. (From
Verworn, after Dreyer.)

The central capsule (rarely inconspicuous and difficult, if not

impossible to demonstrate) is of a substance which resembles chitin,
though its chemical reactions have not been fully studied hitherto,
and indeed vary from species to species. It is composed of a single
layer, except in Phaeodaria, where it is double. The operculum in this
group, i.e. the area around the aperture, is composed of an outer
layer, which is radially thickened, and a thin inner layer; the former is
produced into the projecting tube ("proboscis").

Reproduction in the Radiolaria may be simple fission due to the

binary fission of the nucleus, the capsule, and the ectoplasm in
succession. If this last feature is omitted we have a colonial
organism, composed of the common ectoplasm containing
numerous central capsules; and the genera in which this occurs, all
belonging to the Peripylaea, were formerly separated (as
Polycyttaria) from the remaining Radiolaria (Monocyttaria). They may
either lack a skeleton (Collozoidae, Fig. 22), or have a skeleton of
detached spicules (Sphaerozoidae), or possess latticed shells
(Collosphaeridae) one for each capsule, and would seem therefore
to belong, as only differentiated by their colonial habit, to the several
groups having these respective characters. Fission has been well
studied in Aulacantha (a Phaeodarian) by Borgert.[91] He finds that in
this case the skeleton is divided between the daughter-cells, and the
missing part is regenerated. In cases where this is impossible one of
the daughter-cells retains the old skeleton, and the other escapes as
a bud to form a new skeleton.

Fig. 28.—Shells of Challengeridae: A, Tuscarora; B, Pharyngella; C,

Haeckeliana. (From Wyville Thomson.)

Two modes of reproduction by flagellate zoospores have been

described (Fig. 22). In the one mode all the zoospores are alike—
isospores—and frequently contain a crystal of proteid nature as well
as oil-globules. In the Polycyttaria alone has the second mode of
spore-formation been seen, and that in the same species in which
the formation of isospores occurs. Here "anisospores" are formed,
namely, large "mega-," and small "micro-zoospores." They probably
conjugate as male and female respectively; but neither has the
process been observed, nor has any product of such conjugation
(zygote) been recognised. In every case the formation of the
zoospores only involves the endoplasm: the nucleus first undergoes
brood division, and the plasma within the capsule becomes
concentrated about its offspring, and segregates into the spores; the
extracapsular plasm disintegrates.[92]

The Yellow Cells (Zooxanthella), so frequently found in the

Radiolaria were long thought to be constituents of their body.
Cienkowsky found that when the host died from being kept in
unchanged water, the yellow cells survived and multiplied freely,
often escaping from the gelatinised cell-wall as biflagellate
zoospores. The cell-wall is of cellulose. The cell contains two
chloroplastids, or plates coloured with the vegetal pigment
"diatomin." Besides ordinary transverse fission in the ordinary
encysted state in the ectoplasm of the host, when free they may
pass into what is known as a "Palmella-state," the cell-walls
gelatinising; in this condition they multiply freely, and constitute a
jelly in which the individual cells are seen as rounded bodies. They
contain starch in two forms—large hollow granules, not doubly
refractive, and small solid granules which polarise light. We may
regard them as Chrysomonadaceae (p. 113). Similar organisms
occur in many Anthozoa (see pp. 261, 339, 373 f., 396).
Diatomaceae (yellow Algae with silicified cell-walls) sometimes live
in the jelly of certain Collosphaera. Both these forms live in the state
known as "symbiosis" with their host; i.e. they are in mutually helpful
association, the Radiolarian absorbing salts from the water for the
nutrition of both, and the Alga or Flagellate taking up the CO2 due to
the respiration of the host, and building up organic material, the
surplus of which is doubtless utilised, at least in part, for the nutrition
of the host. A similar union between a Fungus and a coloured
vegetal ("holophytic") organism is known as a Lichen.

The Suctorian Infusorian Amoebophrya is parasitic in the ectoplasm

of certain Acantharia, and in the peculiar genus Sticholonche which
appears to be intermediate between this group and Heliozoa.

The Silicoflagellate family Dictyochidae are found temporarily

embedded in the ectoplasm of some of the Phaeocystina, and have
a skeleton of similar nature. Their true nature was shown by Borgert.

The Amphipod crustacean Hyperia[93] may enter the jelly of the

colonial forms, and feed there at will on the host.[94]

Haeckel, in his Monograph of the Radiolaria of the Challenger

enumerated 739 genera, comprising 4318 species; and Dreyer has
added 6 new genera, comprising 39 species, besides 7 belonging to
known genera. Possibly, as we shall see, many of the species may
be mere states of growth, for it is impossible to study the life-
histories of this group; on the other hand, it is pretty certain that new
forms are likely to be discovered and described. The Radiolaria are
found living at all depths in the sea, by the superficial or deep tow-
net; and some appear to live near the bottom, where the durable
forms of the whole range also settle and accumulate. They thus form
what is known as Radiolarian ooze, which is distinguished from other
shallower deposits chiefly through the disappearance by solution of
all calcareous skeletons, as they slowly fell through the waters
whereon they originally floated at the same time with the siliceous
remains of the Radiolaria. The greatest wealth of forms is found in
tropical seas, though in some places in cold regions large numbers
of individuals of a limited range of species have been found.

Radiolaria of the groups with a pure siliceous skeleton can alone be

fossilised, even the impure siliceous skeleton of the Phaeodaria
readily dissolving in the depths at which they live: they have been
generally described by Ehrenberg's name Polycystineae. Tripolis
(Kieselguhr) of Tertiary ages have been found in many parts of the
globe, consisting largely or mainly of Radiolaria, and representing a
Radiolarian ooze. That of the Miocene of Barbados contains at least
400 species; that of Gruppe at least 130. In Secondary and
Palaeozoic rocks such oozes pass into Radiolarian quartzites (some
as recent as the Jurassic). They occur also in fossilised excrement
(coprolites), and in flint or chert concretions, as far down as the
lowest fossiliferous rocks, the Cambrian. The older forms are simple
Sphaerellaria and Nassellaria. From a synopsis of the history of the
order in Haeckel's Monograph (pp. clxxxvi.-clxxxviii.) we learn that
while a large number of skeletal forms had been described by
Ehrenberg, Huxley in 1851 published the first account of the living
animal. Since then our knowledge has been extended by the labours
of Haeckel, Cienkowsky, R. Hertwig, Karl Brandt, and A. Borgert.

5. Proteomyxa
Sarcodina without a clear ectoplasm, whose active forms are
amoeboid or flagellate, or pass from the latter form to the former;
multiplying chiefly, if not exclusively, by brood-formation in a cyst. No
complete cell-pairing (syngamy) known, though the cytoplasms may
unite into plasmodia; pseudopodia of the amoeboid forms usually
radiate or filose, but without axial filaments. Saprophytic or parasitic
in living animals or plants.

This group is a sort of lumber-room for forms which it is hard to place

under Rhizopoda or Flagellata, and which produce simple cysts for
reproduction, not fructifications like the Mycetozoa. The cyst may be
formed for protection under drought ("hypnocyst"), or as a
preliminary to spore-formation ("sporocyst"). The latter may have a
simple wall (simple sporocyst), or else two or three formed in
succession ("resting cyst"), so as to enable it to resist prolonged
desiccation, etc.: both differing from the hypnocyst in that their
contents undergo brood formation. On encystment any indigestible
food materials are extruded into the cyst, and in the "resting cysts,"
which are usually of at least two layers, this faecal mass lies in the
space between them. The brood-cells escape, either as flagellate-
cells, resembling the simpler Protomastigina, called "flagellulae," and
which often become amoeboid (Fig. 29); or already furnished with
pseudopodia, and called "amoebulae," though they usually recall
Actinophrys rather than Amoeba. In Vampyrella and some others the
amoebulae fuse, and so attain a greater size, which is most probably
advantageous for feeding purposes. But usually it is as a uninucleate
cell that the being encysts. They may feed either by ingestion by the
pseudopodia, by the whole surface contained in a living host-cell, or
by passing a pseudopodium into a host-cell (Fig. 29 5). They may be
divided as follows:—

A. Myxoidea.—Flagella 1-3; zoospores separating at once.

1. Zoosporeae.—Brood-cells escaping as flagellulae, even if they become
amoeboid later. Ciliophrys Cienk.; Pseudospora Cienk. (Fig. 29).

2. Azoosporeae.—Cells never flagellate. Protomyxa Haeckel;

Plasmodiophora Woronin; Vampyrella Cienk.; Serumsporidium L. Pfeiffer.

B. Catallacta.—Brood-cells of cyst on liberation adhering at the centre to

form a spherical colony, multiflagellate; afterwards separating, and becoming
amoeboid. Magosphaera Haeckel (marine).[95]

Fig. 29.—Pseudospora lindstedtii. 1, 2, Flagellate zoospores; 3, young

amoebula, with two contractile vacuoles, one being reconstituted by three
minute formative vacuoles; 4, 5, an amoebula migrating to a fungus hypha
through the wall of which it has sent a long pseudopodium; 6, amoebula full-
grown; 7, 8, mature cells rounded off, protruding a flagellum, before
encysting; 9, young sporocyst; 10, the nucleus has divided into a brood of
eight; 11-14, stages of formation of zoospores. cv, Contractile vacuole; e,
mass of faecal granules; fl, flagellum; n, nucleus, × about 750⁄1.

Plasmodiophora infests the roots of Crucifers, causing the disease

known as "Hanburies," or "fingers and toes," in turnips, etc.
Serumsporidium dwells in the body cavity of small Crustacea. Many
of this group were described by Cienkowsky under the name of
"Monadineae" (in Arch. Mikr. Anat. i. 1865, p. 203). Zopf has added
more than anyone else since then to our knowledge. He
monographed them under Cienkowsky's name, as a subordinate
group of the Myxomycetes, "Pilzthiere oder Schleimpilze," in
Schenk's Handb. d. Bot. vol. iii. pt. ii. (1887). To Lankester (Encycl.
Brit., reprint 1891) we owe the name here adopted. Zopf has
successfully pursued their study in recent papers in his Beitr. Nied.
Org. The Chytridieae, usually ascribed to Fungi, are so closely allied
to this group that Zopf proposes to include at least the Synchytrieae
herein.

This group is very closely allied to Sporozoa; for the absence of

cytogamy, and of sickle-germs,[96] and of the complex spores and
cysts of the Neosporidia, are the only absolute distinctions.

6. Mycetozoa (Myxomycetes, Myxogastres)

Sarcodina moving and feeding by pseudopodia, with no skeleton,
aggregating more or less completely into complex "fructifications"
before forming 1-nucleate resting spores; these may in the first
instance liberate flagellate zoospores, which afterwards become
amoeboid, or may be amoeboid from the first; zoospores capable of
forming hypnocysts from which the contents escape in the original
form.

1. Aggregation taking place without

plastogamy, zoospores amoeboid, with a
clear ectosarc Acrasieae.
Copromyxa Zopf; Dictyostelium Brefeld.
2. Aggregation remaining lax, with merely
thread-like connexions, except when
encystment is to take place; cytoplasm
finely granular throughout; complete
fusion of the cytoplasm doubtful Filoplasmodieae
Labyrinthula Cienk.; Chlamydomyxa Archer; Leydenia (?)
Schaud.
3. Plasmodium formation complete, Myxomycetes.
 eventuating in the formation of a complex
fructification often traversed by elastic,
hygroscopic threads, which by their
contraction scatter the spores; zoospores
usually flagellate at first
Fuligo Hall.; Chondrioderma Rostaf.; Didymium Schrad. (Fig.
30).

I. The Acrasieae are a small group of saprophytes, often in the most

literal sense, though in some cases it has been proved that the
actual food is the bacteria of putrefaction. In them, since no cell-
division takes place in the fructification, it is certain that the
multiplication of the species must be due to the fissions of the
amoeboid zoospores, which often have the habit of Amoeba limax
(Fig. 1, p. 5).

II. Filoplasmodieae.—Chlamydomyxa[97] is a not uncommon

inhabitant of the cells of bog-mosses and bog-pools, and its nutrition
may be holophytic, as it contains chromoplasts; but it can also feed
amoeba-fashion. Labyrinthula is marine, and in its fructification each
of the component cells forms four spores. Leydenia has been found
in the fluid of ascitic dropsy, associated with malignant tumour.

III. Myxomycetes.—The fructification in this group is not formed by

the mere aggregation of the zoospores, but these fuse by their
cytoplasm to form a multinucleate body, the "plasmodium," which,
after moving and growing (with nuclear division) for some time like a
great multinucleate Reticularian, passes into rest, and develops a
fructification by the formation of a complex outer wall; within this the
contents, after multiplication of the nuclei, resolve themselves into
uninucleate spores, each with its own cyst-wall. The fructifications of
this group are often conspicuous, and resemble those of the
Gasteromycetous fungi (e.g., the Puffballs), whence they were at
first called Myxogastres. De Bary first discovered their true nature in
1859, and ever since they have been claimed by botanist and
zoologist alike.

The spore on germination liberates its contents as a minute

flagellate, with a single anterior lash and a contractile vacuole (Fig.
30, C). It soon loses the lash, becomes amoeboid, and feeds on
bacteria, etc. (Fig. 30, D, E). In this state it can pass into hypnocysts,
from which, as from the spores, it emerges as a flagellula. After a
time the amoeboids, which may multiply by fission, fuse on meeting,
so as to form the plasmodium (Fig. 30, F). This contains numerous
nuclei, which multiply as it grows, and numerous contractile
vacuoles. When it attains full size it becomes negatively hydrotactic,
crawls to a dry place, and resolves itself into the fructification. The
external wall, and sometimes a basal support to the fruit, are
differentiated from the outer layer of protoplasm; while the nuclei
within, after undergoing a final bipartition, concentrate each around
an independent portion of plasma, which again is surrounded as a
spore by a cyst-wall. Often the maturing plasmodium within the wall
of the fruit is traversed by a network of anastomosing tubes filled
with liquid, the walls of which become differentiated into membrane
like the fruit-wall, and are continuous therewith. As the fruit ripens
the liquid dries, and the tubes now form a network of hollow threads,
the "capillitium," often with external spiral ridges (Fig. 30, A, B).
These are very hygroscopic, and by their expansion and contraction
determine the rupture of the fruit-wall and the scattering of the
spores.