Energy Transitions and the Future of

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Energy Transitions and
the Future of the
African Energy Sector
Law, Policy and Governance
Edited by Victoria R. Nalule
Energy Transitions and the Future of the African
Energy Sector
 Victoria R. Nalule
Editor

Energy Transitions
and the Future
of the African
Energy Sector
Law, Policy and Governance
Editor
Victoria R. Nalule
School of Social Sciences, CEPMLP
University of Dundee
Dundee, UK

ISBN 978-3-030-56848-1 ISBN 978-3-030-56849-8 (eBook)

https://doi.org/10.1007/978-3-030-56849-8

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To the African Youth, may you be inspired by this book to actively take part
in shaping the future of the African energy sector
 Acknowledgements

I want to take this opportunity to thank all those who contributed

chapters to this book. A special thanks go out to Ayebare Rukundo
Tom for coming up with the brilliant idea of having an edited
African book project featuring different experts. I also do sincerely
thank Noreen Kidunduhu and Hendrica Rabophala for their editorial
assistance. I again sincerely thank Rachael Ballard and Joanna O’Neil
for their help in getting the work published on time. The African
Energy and Minerals Management Initiative (AEMI) was also instru-
mental in bringing together different African experts (www.afrienergymi
nerals.org).

vii
 Contents

Part I Introduction

1 How to Respond to Energy Transitions in Africa:

Introducing the Energy Progression Dialogue 3
Victoria R. Nalule
1.1 Introduction: Aim of the Book 3
1.2 The Future and Role of Fossil Fuels in Africa
in the Energy Transition Era 7
1.3 Introducing the Energy Progression Dialogue:
Energy Progression V Energy Transitions 14
1.4 Energy Transition Indicators in African
Countries: Legal Analysis 23
1.5 Outline of the Book 29
References 33

ix
x Contents

2 African Energy Challenges in the Transition Era: The

Role of Regional Cooperation 37
Macdonald Irowarisima
2.1 Introduction 37
2.2 Overview of the Character of Energy Resource
in Sub-Saharan Africa Through the Lens
of Ecowas and SADC Regions 41
2.3 Current Energy Challenges Applicable
to ECOWAS and SADC 49
2.4 Areas of Regional Cooperative Role in Tackling
the Energy Challenge 58
2.5 Conclusion/Recommendation 67

3 Energy Transition in Africa: Context, Barriers

and Strategies 73
Noreen Kidunduhu
3.1 Introduction 73
3.2 Energy Landscape in Africa 78
3.3 Barriers to the Energy Transition 85
3.4 Countermeasures and Suggestions 95
3.5 Conclusion 104
Bibliography 106

4 Nuclear Energy & Energy Transitions: Prospects,

Challenges and Safeguards in Sub-Saharan Africa 113
Susan Nakanwagi
4.1 Introduction 113
4.2 Nuclear Energy: Access to Energy and Climate
Change 116
4.3 The International Legal Order on Nuclear Energy 122
4.4 The Development and Regulation of Nuclear
Energy in Sub-Saharan Africa: Lessons
from the EU and France 127
4.5 Recommendations and Conclusions 131
Bibliography 133
 Contents xi

Part II Geographies of Energy Transition: Managing the

Decline of Fossil Fuels in Africa

5 Role of Law in the Energy Transitions in Africa:

Case Study of Nigeria’s Electricity Laws and Off-Grid
Renewable Energy Development 141
Michael Uche Ukponu, Yusuf Sulayman, and Kester Oyibo
5.1 Introduction 141
5.2 Overview of Nigeria’s Renewable Energy
Regulatory Framework 146
5.3 A Critical Analysis of the Conflict of Electricity
Laws 151
5.4 Contemporary Approaches to Actualizing Rapid
Renewable Energy Development from Legal
and Governance Perspectives 165
5.5 Conclusion 178
Bibliography 180

6 The Status and Future of Charcoal in the Energy

Transition Era in Sub-Saharan Africa: Observations
from Uganda 189
Catherine Nabukalu and Reto Gieré
6.1 Introduction 189
6.2 Environmental and Health Implications
of Charcoal Production and Consumption 191
6.3 Methodology 194
6.4 The Status of Raw Material Procurement
for Charcoal Production in Africa 195
6.5 Charcoal in the Final Market 203
6.6 Optionality in the Energy Mix: When
Consumers Choose Charcoal Over Modern
Alternatives 209
6.7 Beyond Market Dynamics: Infrastructural
Barriers to the Consumption of Modern Energy
Alternatives in Africa 212
xii Contents

6.8 Charcoal in International Markets and Africa’s

Role as a Major Exporter 215
6.9 Summary and Conclusions 219
References 222

7 Will a Transition to Renewable Energy Promote

Energy Security Amid Energy Crisis in Nigeria? 231
Cosmos Nike Nwedu
7.1 Introduction 231
7.2 Energy Security in Energy Transition Regime:
The Case of Nigeria 233
7.3 Forms and Prospects of Renewables for Energy
Security in Nigeria 238
7.4 Overcoming Renewable Energy Development
Challenges in Nigeria 251
7.5 Conclusion 257
Bibliography 258

8 Renewable Energy Development in Egypt

and Transitioning to a Low-Carbon Economy 265
Mostafa Elshazly
8.1 Introduction 265
8.2 The Institutional Structure of the Egyptian
Energy Sector 268
8.3 Legal Measures and Policies in Place that Address
the Promotion and Management of Renewable
Forms of Energy and Energy Efficiency 271
8.4 Progress of Renewable Energy Projects in Egypt 277
8.5 Challenges Hindering Renewable Energy Projects
in Egypt 279
8.6 Proposed Measures to Encourage Players
in the Hydrocarbon Sector to Reduce Their
Carbon Footprint/Emissions 282
8.7 Conclusion 284
References 285
 Contents xiii

9 Transitioning to a Low-Carbon Economy

and Renewable Energy Developments in Uganda:
Challenges and Opportunities for Small-Scale
Renewable Energy 287
Marvin Tumusiime
9.1 Introduction 287
9.2 Main Constraints to Uganda’s Efforts
to Transition to a Low-Carbon Economy 289
9.3 How Can Uganda Effectively Transition
to a Low-Carbon Economy? 292
9.4 What Are the Transition Needs for Small-Scale
Renewable Energy? 299
9.5 Conclusion 303
References 304

Part III Extractives in the Energy Transition Era: Key

Concepts

10 Local Content Policies in the Energy Transition

Era in Africa: A Case Study of the East African Oil
and Gas Industry 311
Rukonge S. Muhongo
10.1 Introduction 311
10.2 Breaking the Enclave in the East African Oil
and Gas Industry 315
10.3 Factors Affecting Local Content Design in East
Africa 318
10.4 The Advantages and Disadvantages of Local
Content Policies 323
10.5 The Norwegian and Nigerian Case Study:
Lessons to Be Drawn for Emerging
Resource-Rich Countries 326
10.6 Analysis of the Norwegian and Brazilian Local
Content Regimes 329
xiv Contents

10.7 Lessons for Emerging Oil and Gas Producers

Such as Kenya, Tanzania and Uganda 331
10.8 Creating a Regional Content Policy for the Oil
and Gas Industry in East Africa 333
10.9 Conclusion 335
Bibliography 337

11 Social Licence to Operate in the Energy Transition

Era: Case Study of the East African Oil and Gas Sector 341
Wairimu Karanja and Nduta Njenga
11.1 Introduction to Social Licence to Operate 341
11.2 The State of Oil and Gas and SLO in East Africa 347
11.3 East African Oil and Gas Laws in Relation
to SLO 353
11.4 Corporate and Government Initiatives to Seek
SLO in East Africa 360
11.5 Conclusion 365
Bibliography 366

12 Gender Justice in the Energy Transition Era:

Exploring Gender and Technology in the Extractives
Sector 371
Alaka Lugonzo and Kennedy Chege
12.1 Introduction 371
12.2 Why It Is Imperative to Include Women
in the Global Economy 375
12.3 The Need for Automation in the Extractives
Industries 378
12.4 Sector Outlook: The Position of Women
in the Extractives Sector 380
12.5 Artificial Intelligence and the Future
of Extractives 384
12.6 Proposed Solutions to the Challenge of Gender
Inclusivity in the Extractives Sectors 385
12.7 Conclusion 391
Bibliography 393
 Contents xv

13 A Critical Intertemporal Analysis of Uganda’s Fiscal

Regime in the Energy Transition Era: Are Uganda’s
Upstream Projects Still Viable? 397
Ayebare Rukundo Tom
13.1 Introduction 397
13.2 Literature Review 399
13.3 Methodology—Evaluation of Economics
of Fiscal Terms and Alternative Regime 406
13.4 Results and Analysis 409
13.5 Conclusion 424
References 425

14 Energy Transitions and Environmental Protection:

Environmental Impact of the Oil and Gas Production
Activities in South Sudan 429
Peter Reat Gatkuoth
14.1 Introduction 429
14.2 Environmental Impacts of Oil and Gas in South
Sudan 431
14.3 Legal Framework Regulating the Oil and Gas
Impacts on the Environment 437
14.4 Conclusion 446
Bibliography 447

15 Steadfast for East Africa: Powering Energy Dreams

Through Regional Cooperation 449
Japhet Miano Kariuki
15.1 Introduction 449
15.2 Energy Investments in East Africa: Issues
of Concern 451
15.3 Integrated Feed-in Tariff Structure for the East
African Community: Lessons from Thailand 455
15.4 Conclusion 462
References 464
xvi Contents

Part IV Epilogue

16 Extractives and Beyond: Managing the Energy

Transition in Africa 469
Victoria R. Nalule

Index 473
 Notes on Contributors

Kennedy Chege (Kenyan national) is Ph.D. Law student at the Univer-

sity of Cape Town (UCT) in South Africa, and a Researcher at the
NRF/DST SARChI: Mineral Law in Africa (MLiA) Research Chair
at UCT. His Doctoral research relates to mining law, specifically Oil
and Gas Law, incorporating aspects of Competition/Antitrust Law.
Kennedy is also a practitioner in Africa’s largest law firm, ENSafrica,
dealing with matters relating to Mineral & Mining Law, Energy Law,
Competition/Antitrust Law and general corporate and commercial work.
Kennedy also holds a Master of Laws (LL.M.) Degree in Commercial
Law, coupled with Mining Law; a Bachelor of Laws (LL.B.) Degree, and
a Bachelor of Social Sciences (BSocSc) Degree majoring in Public Policy
and Administration, all from UCT. Kennedy’s interests transverse across
Mining and Mineral Law, Oil and Gas/Petroleum Law, Energy Law,
Energy Justice, Natural Resources Law, Resource Revenue Law/Taxation,
Dispute Resolution and related fields.
Dr. Mostafa Elshazly has diverse experience in legal aspects of Energy,
Mining, Oil and Gas (upstream and downstream) projects. He has

xvii
xviii Notes on Contributors

recently advised several International Oil Companies and other compa-

nies in different areas, furthermore he assists governmental bodies in
policymaking process, management and development of their natural
resources.
Before Joining ZH&P, Dr. Mostafa Elshazly worked in several places
including the Egyptian Ministry of Petroleum and The Egyptian General
Petroleum Corporation (EGPC) among others.
Dr. Mostafa holds a Ph.D. in Commercial Law from Cairo University,
he achieved his LL.B. from Faculty of Law (English Section), Alexan-
dria University in 2007, he is a part-time lecturer at several Egyptian
universities including Cairo University and he is an acting member of the
Association of International Petroleum Negotiators (AIPN). Dr. Elshazly
is the author of several articles and papers published by highly ranked
international journals.
Peter Reat Gatkuoth is currently a Director of General Services in
Nile Drilling and Services under the Minister of Petroleum in South
Sudan. Gatkuoth has served in various departments in Dar Petroleum
Operating Company that include Section Head for Policy and Services,
Section Head of Material Management, Director of Technical Services
and Director of General Services in Dar Petroleum until October 2019.
Prior to the abovementioned positions, Gatkuoth served in various
organizations and institutions in Western and Central Canada before
returning back to South Sudan. His positions away from home included
Settlement Counselor in Catholic Social Services (Edmonton), Employ-
ment Counselor in Open Door Society and West Care Facility Services
Branch Manager in the City of Regina and Saskatoon. He has a B.A.
in Sociology (Concordia University of Edmonton), M.A., Int’l Law and
Human Rights (UN Mandated University for Peace), LL.M. Oil and Gas
(Uganda Christian University) and he is currently pursuing a Master of
Public Policy in University of Juba.
Professor Reto Gieré is a Professor and the Chair of Earth and Envi-
ronmental Science programmes at the University of Pennsylvania. He
received his Ph.D. from ETH Zürich, Switzerland. He has held appoint-
ments at Purdue University, the Australian Nuclear Science & Tech-
nology Organization, Albert-Ludwigs-Universität, and the Carnegie
 Notes on Contributors xix

Institution for Science, among others. His broad research interests

involve environmental geochemistry, energy and waste, mineralogy and
petrology, and health impacts of atmospheric pollution. Reto is a
member of the University of Pennsylvania’s Centre for Excellence in
Environmental Toxicology (CEET). Additionally, Reto is an Editor of
the Journal of Petrology and the Chief Editor of the European Journal
of Mineralogy. He has supervised theses of 21 Ph.D. and 36 Masters
students.
Macdonald Irowarisima, Ph.D. is a Nigerian, born in Rivers State of
Nigeria in the early 1980s. Began his academic career in Nigeria. He
obtained his first school leaving certificate from Federal Government
College Kwali Abuja Nigeria. He later did a short certificate programme
at the College of Arts and Science, Aguda-Ama Epie, in Bayelsa State-
Nigeria. In 2015, he began his legal career studies at the Prestigious
Ahmadu Bello University Zaria, Nigeria where he graduated in 2008.
As part of the continued legal education, he enrolled at the Nigerian Law
School, Enugu State Chapter in Nigeria between 2008 and 2009. He did
his National Youth Service Corps programme in 2010 at the Petroleum
Technology Development Fund (PTDF) at the legal department of the
Fund. Afterwards, he decided to pursue an LL.M. master’s programme in
Energy Law and Policy between 2011 and 2012 at the Centre for Energy,
Mineral Law and Policy, University of Dundee, United Kingdom. The
author further proceeded and concluded a Doctorate degree in Energy
Law and Environmental Sustainability at the same University. The author
has a had few years of experience in legal practice in Nigeria and teaching
experiences. He was the Head of legal research team while undergoing
his legal practice at the Bola Aidi & Co Solicitors in Nigeria. Since
then, the author has been active in both collaborative research projects
like the United Nations project in conjunction with the Centre for
Energy Mineral Law and Law Policy at the University of Dundee on
how to assist nations transit from fossil fuel to renewable sources of
energy as a means of achieving energy security and efficiency. Macdonald
Irowarisima has co-authored an article titled ‘Identifying Policy and Legal
Issues for Shale Gas development in Algeria: A SWOT Analysis’. He is
xx Notes on Contributors

passionate in setting policy directions in dealing with complex energy-

related issues and formulating regulatory frameworks for the governance
of the operations of the energy sector globally.
Ms. Wairimu Karanja is Director, Wairimu & Co, a specialist legal
advisory firm focusing on corporate law, energy and natural resources
(ENR), investment and trade matters in Kenya and internationally. She
has 12 years’ experience and previously worked in leading Kenyan and
international firms. Wairimu advised on ENR project contracting and
finance, commercial transactions, immigration and employment and
private equity. She is recognized by Who’s Who Legal 2019 as a future
leader in international arbitration. Wairimu holds a Master’s Degree in
Energy Law and Policy. She is also the University of Dundee’s Extractives
Hub Representative for Kenya. LinkedIn: https://www.linkedin.com/in/
wairimu-karanja-06680861/.
Japhet Miano Kariuki was born in Nairobi, Kenya, he has had a
diverse and varied career, spending much of his subsequent career
working and studying abroad in Europe, America and now in Asia.
Currently, with the World Energy Council’s Scenario Study Group.
Prior to that, he worked in the extractive industry with OLLI Consulting
Group Inc. (Philippines), oil and gas for Shell (Philippines), logistics
for Deutsche Post DHL (Germany), financial services for BNP Paribas
Fortis Investment Bank (Belgium), Citibank (Kenya) and management
consulting for Spatial Systems Architects (Kenya).
He holds a Master’s Degree in financial management from the Vlerick
Business School (Belgium).
Noreen Kidunduhu is a Kenya qualified lawyer whose practice focuses
on international energy law and particularly providing dispute, corpo-
rate and project advisory services as well as practical legal, technical
and commercial solutions to governments, energy and mining compa-
nies, investors and financial institutions in the Eastern Africa region. She
holds a Master’s Degree in International Energy Law and Policy from the
Centre for Energy Petroleum and Mineral Law and Policy (CEPMLP)
at the University of Dundee. She is also a Regional Representative of
 Notes on Contributors xxi

London Court of International’s Arbitration’s Young International Arbi-

trator’s Group and sits on the Law Society of Kenya’s Alternative Dispute
Resolution Committee.
Alaka Lugonzo is an Extractives Sector Consultant with a legal back-
ground and a specialization in Energy, Oil Gas and Mining aspects of the
extractives industry. She is an Advocate of the High Court of Kenya and
has an (LL.M.) in Energy Law and Policy from the University of Dundee
in Scotland. Her core interest and passion is in Policy and Governance,
Local Content, Technology and Gender in the Extractives sector. Alaka’s
experience includes growing new organizations through networking,
fundraising and cultivating relationships, writing and research, teaching
and training, speaking and moderating in conferences, corporate gover-
nance, policy advocacy and capacity building. Alaka is the recipient of the
Women in Business Award 2019 under the Environmental and Natural
Resources Category.
Dr. Rukonge S. Muhongo is an oil, gas and energy law consultant and
lecturer. Currently Dr. Muhongo is a visiting lecturer at the University
of Eduardo Mondlane in Maputo, Mozambique and a Research Assis-
tant for Professor Raphael Heffron in the Solar Justice Project at the
University of Dundee. A contributor to the Energy Justice discourse
with notable contributions in key journals such as Extractive Industry and
Society, and the Global Energy law and Sustainability Journal of the Centre
for Energy, Petroleum and Mineral Law and Policy.
Dr. Muhongo just completed his Ph.D. in the Development of Local
Content Policies, using Energy Justice as an analytical framework from
the University of Dundee. He holds a B.A.(Crim) and LLB degrees both
from the University of Pretoria, South Africa as well as an LLM in Oil
Gas Law and Policy from the University of Dundee, United Kingdom.
Catherine Nabukalu is a graduate of the Master of Environmental
Studies at the University of Pennsylvania (US) and B.Sc. in Logistics
and Supply Chain Management at Aston University (UK). Her research
focuses on secondary energy and the global charcoal supply chain.
Her main question is whether international energy markets have indeed
responded positively to traditional solutions, such as more electrification.
xxii Notes on Contributors

The research explores why charcoal is still a popular energy alternative

even where grid connections are prevalent because charcoal production
has several implications, mainly energy-related forest loss, acute in the
Sub-Saharan region. Furthermore, she examines the status of waste
management along this value chain, as well as public health implications
from charcoal consumption. Currently, she is a Project Coordinator
for Energy Efficiency in Washington DC, at the District of Columbia’s
Sustainable Energy Utility (DCSEU).
Ms. Susan Nakanwagi is a lawyer of the High Court of Uganda,
and a member of the Uganda and East African Law Societies. She is
also working with the International Energy Agency (IEA), in Paris,
France where she makes a comparable analysis on policy and regulatory
approaches to reducing methane emissions. Susan is also the Assistant
Executive Director of the African Energy and Mining Management
Initiative (AEMI) where she also heads the Women in Energy and
Mining Empowerment Program (WEM).
Ms. Susan is currently a Ph.D. scholar at the Centre for Energy,
Petroleum, Mineral Law and Policy—University of Dundee, United
Kingdom. She also holds an LL.M. in Natural Resource Law and Policy
(distinction)—the University of Dundee, a Post Graduate Diploma in
Legal Practice from Law Development Centre, Uganda and a Bachelor
of Laws (Honours) from Makerere University, Uganda. She further has
a Certificate of Distinction in ‘Natural Resources for Sustainable Devel-
opment: The Fundamentals of Oil, Gas and Mining Governance’, from
the SDG Academy.
Dr. Victoria R. Nalule is an Energy and Mining professional & consul-
tant with extensive experience working on various projects in the
different parts of the Globe. She is a holder of a Ph.D. in International
Energy Law and Policy. Victoria is the Founder and Executive Director
of the African Energy and Minerals Management Initiative (AEMI).
She is currently involved as a Research Fellow with the DFID-funded,
Extractives Hub project, based at CEPMLP, University of Dundee, UK.
Victoria is an author and has widely published on topics relating to oil,
gas, renewable energy, climate change and mining in Africa. Her latest
three books being, Mining and the Law in Africa: Exploring the Social and
 Notes on Contributors xxiii

Environmental Impacts; Energy Transitions and the Future of the African

Energy Sector: Law, Policy and Governance; and another book on Energy
Poverty and Access Challenges in Sub-saharan Africa: The Role of Region-
alism. She also has an upcoming book with Hart publishers focused on
Land Law and Extractives.
Victoria offers extensive experience in the Energy and Mining sectors
having worked with various institutions; regional and international
organizations including assignments for The Queen Mary University
of London (EU Energy Project); The International Energy Charter
Secretariat in Belgium; The Columbia Center on Sustainable Invest-
ment in New York; the East African Community Secretariat (EAC) in
Tanzania; the University of Dundee in Scotland; the Southern African
Development Community Secretariat (SADC) in Botswana; Interna-
tional Arbitration Case Law in New York (editor); Institute of Petroleum
Studies-Kampala; Uganda Christian University; Journal of Mines, Metals
and Fuels (Editorial Board Member). She has also worked with both
the private and public legal sectors of Uganda including Kakuru & Co.
Advocates and the Anti-Corruption Court of Uganda.
She has presented as a speaker and panellist in several forums and
conferences discussing topics concerning oil and gas; climate change,
renewable energy, mining, energy poverty and access to mention but a
few. She has also advised African governments on oil, gas and mining
projects including appearing as an expert witness before the Commission
of Inquiry into Land matters in Uganda; presenting comments on the
South African Petroleum Bill before the country’s Policy Makers.
Victoria has an active YouTube Channel, i.e. Victoria Nalule and
Podcast, i.e. Dr. Victoria Nalule, both focused on energy and mining
discussions. She is one of the few people who got her Ph.D. in less than
3 years below the age of 30 years.
Ms. Nduta Njenga is an Oxford Policy Fellow (Energy Advisor) at the
Ministry of Water, Irrigation & Energy (Ethiopia), Consultant at W &
Co. (Kenya) and Director at the African Energy and Minerals Manage-
ment Initiative (AEMI, Uganda). She is a practicing Advocate with 8+
years’ experience working in the Public Sector in the areas of Resource
xxiv Notes on Contributors

Governance, Energy and Extractives Management. Nduta holds a Post-

graduate LL.M. in Energy Law and Policy from CEPMLP, UK and is
a featured author in the Oil, Gas and Energy Law (OGEL) Journal , the
Palgrave Handbook of Managing Fossil Fuels and Energy Transitions and
the Meteor Springer Encyclopaedia of Mineral and Energy Policy.
Cosmos Nike Nwedu is a solicitor and advocate of the Supreme Court
of Nigeria, and currently in private legal practice with the law firm of
U.M. Usulor & Associates. He also undertakes research and teaching
activities at the Faculty of Law, Federal University Ndufu-Alike Ikwo,
Nigeria. He received his LL.M. degree in Climate Change and Energy
Law and Policy from CEPMLP, University of Dundee, UK, B.L from
Nigerian Law School, Abuja and LLB Hons degree from Ebonyi State
University, Nigeria. His research interest is in the broad areas of climate
change and energy law and policy, clinical legal education and interna-
tional arbitration.
Kester Oyibo, AMEI, MCIArb is a licensed Nigerian legal practitioner.
He has over seven years’ experience in the delivery of legal services
to individuals and corporate entities in the areas of dispute resolution
and energy transactions. He holds a Bachelor of Laws from Madonna
University (Nigeria) and a Master of Laws from the Centre for Energy,
Petroleum and Mineral Law and Policy, University of Dundee (UK) with
specialization in International Energy Law and Policy. He is a member
of several professional international organizations including the Energy
Institute, Association of International Petroleum Negotiators, Society of
Petroleum Engineers and the Chartered Institute of Arbitrators, UK.
Kester is a Senior Associate in Compos Mentis, a boutique law firm
in Nigeria. He is also a volunteer for the UK Department of Interna-
tional Development (DFID) funded Extractive HUB project and partic-
ipated in the Energy law legislative draft project of the Centre for Energy,
Petroleum and Mineral Law and Policy, University of Dundee. Kester is
a Fellow of the United States’ YALI RLC Emerging Leaders Programme,
West Africa, and continues to make valid contributions to social justice
and energy Policy discourses in Nigeria.
 Notes on Contributors xxv

Yusuf Sulayman, Esq., LL.M. a Partner in the law-firm of Adingwu,

Maude & Sulayman LP and a Legal Consultant for One Community
Inc., (USA), is a research enthusiast and passionate social entrepreneur.
With One Community Inc., Yusuf has drafted numerous high-level
agreements while being heavily involved in other academic research
works as a research consultant.
From 2017 to 2019, he was a fellow of Teach For Nigeria, an initia-
tive for helping primary school children from underserved communi-
ties in Nigeria in the areas of literacy, numeracy and science, while also
ideating innovative projects like ArtScholar and ToiletUp Schools to
confront educational inequities in Nigeria. Yusuf was recently appointed
as a member of the Board of a non-profit organization—Advocates
against Inequities Initiative—where he is helping to design and imple-
ment strategies to advocate and take actions against social injustices in
Nigeria.
Mr. Ayebare Rukundo Tom is a Petroleum economist with over seven
years’ experience in cost, financial and economic aspects of Uganda’s
petroleum and development sectors executed in Uganda, France and the
United Kingdom. Currently, he works with the Petroleum Authority of
Uganda as the Manager Economic and Financial analysis. Previously, he
worked with the Ministry of Energy and Mineral Development in roles
relating to the economic and commercial regulation of the oil and gas
sector. He has also worked with National Housing and Construction
company and Uganda National Roads Authority in various capacities.
He possesses a Master of Science in Oil and Gas Economics from the
University of Dundee and a Bachelor’s Degree in Quantity Surveying
from Makerere University.
Marvin Tumusiime is an energy economist and consultant currently
serving as the Programme Manager in the Ugandan chapter of New
Energy Nexus, an international organization that supports clean energy
entrepreneurs and its headquarters are located in California with offices
in Singapore, China, Philippines, Indonesia, India and East Africa.
The Ugandan chapter now supports 97 clean energy enterprises with
financing, reaching 87,000 people. He has project-managed on different
energy access projects for Mercy Corps, Response Innovation Lab and
xxvi Notes on Contributors

more. He is a member and contributor with the Affordable Energy for

Humanity and a Business Development Support advisor for SEED. Aside
from that, he has written blog articles on clean energy uptake in Sub-
Saharan Africa. Marvin graduated with a Master of Science in Energy
Studies from the University of Dundee in 2016 and prior to that, grad-
uated with a B.A. (Hons) in Finance awarded by the University of East
London in 2013.
Michael Uche Ukponu is a Barrister and Solicitor of the Supreme
Court of Nigeria. He is currently a Senior Associate at The Law Part-
ners (Barristers and Solicitors), a reputable law firm with its headquarters
in Abuja, Nigeria, specializing in energy and environmental law. It was
on the strength of his passion for sustainable development in the energy,
natural resources and the environment sectors that he was granted the
Australia Awards Scholarship by the Australian Government to under-
take studies in Energy and Resources Law (LL.M.) at The University of
Melbourne, Australia. He is a member of notable international profes-
sional associations, namely the Association of International Petroleum
Negotiators; the Chartered Institute of Arbitrators, UK; and the Envi-
ronment Institute of Australia and New Zealand and has published
research articles related to renewable energy regulation and access to
environmental justice.
 Abbreviations

AETR Average Effective Tax Rate

BAU Business As Usual
BEP Break-Even Price
BITs Bilateral Investment Treaties
BOO Build- Own- Operate
BOPD Barrels of Oil Per Day
BSC Brussels Supplementary Convention
BSCF Billions of Standard Cubic Feet of Gas
CAPEX Capital Expenditure
CCS Carbon Capture Storage
CDM Clean Development Mechanism
CES Conventional Energy Sources
CFRN Constitution of the Federal Republic of Nigeria
COAG Council of Australian Government
COP Conference of the Parties
COVID-19 Coronavirus Disease 2019
CPF Central Processing Facility
E&P Exploration and Production
EBRD European Bank for Reconstruction and Development
EC European Community

xxvii
xxviii Abbreviations

EDB Ease of Doing Business

EEHC Egyptian Electricity Holding Company
EETC Egyptian Electricity Transmission Company
EG Embedded Generation
EGAS Egyptian Natural Gas Holding Company
EGPC The Egyptian General Petroleum Corporation
EgyptERA Egyptian Electric Utility and Consumer Protection Regulatory
Agency
EIA Energy Information Administration
EPA Environmental Protection Agency
EPC Engineering and Procurement Contract
EPS Embedded Power Scheme
EPSRA Electric Power Sector Reform Act
ERGP Economic Recovery and Growth Plan
EU European Union
FIPs Feed-in-Premiums
FITs Feed-in-Tariffs
FY Financial Year
GHG Green House Gas
GHI Global Horizon Irradiation
GW Gigawatt
GWh Gigawatt hour
IAEA International Atomic Energy Agency
IANEC Inter-American Nuclear Energy Commission
ICRP International Commission on Radiological Protection
IEA International Energy Agency
IEDN Independent Electricity Distribution Network
IET International Emissions Trading
IHA International Hydropower Association
INDC Intended Nationally Determined Contribution
INES International Nuclear Event Scale
IOCs International Oil Companies
IPCC Intergovernmental Panel on Climate Change
IRENA International Renewable Energy Agency
ISES Integrated Sustainable Energy Strategy
JI Joint Implementation
LEPSRL Lagos State Electric Power Sector Reform Law
LESB Lagos State Electricity Board
LGA Local Government Area
 Abbreviations xxix

LNG Liquefied Natural Gas

M/S Meter per Second
M2 Square Metre
MJ Megajoule
MOERE Ministry of Electricity and Renewable Energy
MOP Ministry of Petroleum and Mineral Resources
MTOE Million Tonnes of Oil Equivalent
MW Megawatt
NACOP Nigeria National Council on Power
NDCs Nationally Determined Contributions
NDPHC Niger Delta Power Holding Company Limited
NERC Nigerian Electricity Regulatory Commission
NESI Nigerian Electricity Supply Industry
NESP Nigerian Energy Support Programme
NIPP National Integrated Power Project
NIREDA National Renewable Energy Development Agency
NREA New and Renewable energy Authority
NREAP National Renewable Energy Action Plans
NREEEP National Renewable Energy and Energy Efficiency Policy
OECD Organization for Economic Co-operation and Development
PC Paris Convention
PCCA Paris Climate Change Agreement
PPA Power Purchase Agreement
PSAs Production Sharing Agreements
PSRP Power Sector Recovery Programme
RE Renewable Energy
REA Rural Electrification Agency
REF Rural Electrification Fund
REL Renewable Energy Law
REMP Renewable Energy Master Plan
RES Renewable Energy Sources
RESIP Rural Electrification Strategy and Implementation Plan
RET Renewable Energy Target
SDGs Sustainable Development Goals
SDR Special Drawing Rights
SE4ALL Sustainable Energy for All
SEC Supreme Energy Council
SSA Sub-Saharan-Africa
TCF Trillion Cubic Feet
xxx Abbreviations

TEPCO Tokyo Electric Power Company

TPES Total Primary Energy Supply
TWh Terawatt-hours
UN United Nations
UNCED United Nations Conference on Environment and Development
UNIDO United Nations Industrial Development Organization
USA United States of America
USAID United States Agency for International Development
WTE Waste-to-Energy
 List of Figures

Fig. 5.1 A mnemonical illustration of the conflicts existing

within Nigeria’s electricity regulatory framework 153
Fig. 6.1 Satellite image showing the location of Uganda’s
capital Kampala and of other major towns as well
as the districts mentioned in the text (labelled in
pink) and/or some of the sites investigated during the
field work. Image from Google Earth 2018 (Image
Landsat/Copernicus). DRC = Democratic Republic
of the Congo 193
Fig. 6.2 Rudimentary charcoal production in Uganda. a A
hand tool (panga) used in manual tree-felling; b A pile
of logs ready to be covered with damp soil; c Active
pyrolysis process showing the release of moisture from
the earth-mound kiln; d Tree regeneration in the
open forest near a charcoal production site (a–d were
taken near Naminato bridge in the Nwoya district);
e Manual harvesting using a rake near Kikumbi,
Mityana district; f Harvesting with bare hands while
packaging near Bulera, Mityana district 196

xxxi
Another random document with
no related content on Scribd:
 This sub-order has been established for the reception of the
curious genus Opilioacarus.
Fam. Opilioacaridae.—Mites with segmented abdomen, leg-like
palps, chelate chelicerae, and two pairs of eyes. There are four dorsal
abdominal stigmata. Four species of the sole genus Opilioacarus
have been recorded, O. segmentatus from Algeria, O. italicus from
Italy, O. arabicus from Arabia, and O. platensis[373] from South
America.
 APPENDICES TO ARACHNIDA
I. and II.

TARDIGRADA AND PENTASTOMIDA

BY

ARTHUR E. SHIPLEY, M.A., F.R.S.

Fellow and Tutor of Christ’s College, Cambridge, and Reader in

Zoology in the University
 CHAPTER XIX
TARDIGRADA

OCCURRENCE—ECDYSIS—STRUCTURE—DEVELOPMENT
—AFFINITIES—BIOLOGY—DESICCATION—PARASITES—
SYSTEMATIC

The animals dealt with in this chapter lead obscure lives, remote
from the world, and few but the specialist have any first-hand
acquaintance with them. Structurally they are thought to show
affinities with the Arachnida, but their connexion with this Phylum is
at best a remote one.
Tardigrades are amongst the most minute multicellular animals
which exist, and their small size—averaging from ⅓ to 1 mm. in
length—and retiring habits render them very inconspicuous, so that
as a rule they are overlooked; yet Max Schultze[374] asserts that
without any doubt they are the most widely distributed of all
segmented animals. They are found amongst moss, etc., growing in
gutters, on roofs, trees or in ditches, and in such numbers that
Schultze states that almost any piece of moss the size of a pea will, if
closely examined, yield some members of this group, but they are
very difficult to see. The genus Macrobiotus especially affects the
roots of moss growing on stones and old walls. M. macronyx lives
entirely in fresh water, and Lydella dujardini and Echiniscoides
sigismundi are marine; all other species are practically terrestrial,
though inhabiting very damp places.
In searching amongst the heather of the Scotch moors for the ova
and embryos of the Nematodes which infest the alimentary canal of
the grouse, I have recently adopted a method not, as far as I am
aware, in use before, and one which in every case has yielded a good
supply of Tardigrades otherwise so difficult to find. The method is to
soak the heather in water for some hours and then thoroughly shake
it, or to shake it gently in a rocking machine for some hours. The
sediment is allowed to settle, and is then removed with a pipette and
placed in a centrifugaliser. A few turns of the handle are sufficient to
concentrate at the bottom of the test-tubes a perfectly amazing
amount of cryptozoic animal life, and amongst other forms I have
never failed to find Tardigrades.
Many Tardigrades are very
transparent; their cells are large,
and arranged in a beautifully
symmetrical manner; and since
those of them that live in moss,
and at times undergo desiccation,
are readily thrown into a perfectly
motionless state, during which
they may be examined at leisure,
it is not surprising that these little
creatures have been a favourite
object for histological research.
One way to produce the above-
mentioned stillness is partly to
asphyxiate the animals by placing
them in water which has been
boiled, and covering the surface
of the water with a film of oil.
The whole body is enclosed in a
thin transparent cuticle, which
must be pierced by a needle if it
be desired to stain the tissues of
the interior. As a rule the cuticle
is of the same thickness all over
the body, but in the genus
Fig. 249.—Dorsal view of Echiniscus
Echiniscus the cuticle of the
testudo, C. Sch., × 200, showing the
four segments 1, 2, 3, 4. (From Doyère.) dorsal surface is arranged in
thickened plates, and these plates
are finely granulated. From time
to time the cuticle is cast, and this is a lengthy process, so that it is
not unusual to find a Tardigrade ensheathed in two cuticles, the
outer of which is being rubbed off. The Macrobioti lay their eggs in
their cast cuticle (Fig. 250). The end of each of the eight legs bears
forked claws of cuticular origin. The legs are not jointed except in the
genus Lydella, where two divisions are apparent.
 Within the cuticle is the
epidermis, a single layer of cells
arranged in regular longitudinal
and transverse rows along the
upper and under surface, where
the cells are as uniformly
arranged and as rectangular as
bricks. The cells on the sides of
the body are polygonal, and not in
such definite rows. The nuclei
show the same diagrammatic
symmetry as the cells which
contain them, and lie in the same
relative position in neighbouring
cells. In a few places, such as the
end of each limb and around the
mouth and arms, the cells of the
epidermis are heaped up and
form a clump or ridge. In some
genera a deposit of pigment in the
epidermis, which increases as the
animal grows old, obscures the
internal structures. It is generally
brown, black, or red in colour.
The cuticle and epidermis
enclose a space in which the
various internal organs lie. This
space is traversed by numerous
symmetrically disposed muscle- Fig. 250.—Cast-off cuticle of
fibres, and contains a clear fluid— Macrobiotus tetradactylus, Gr., ×
the blood—which everywhere about 150, containing four eggs in
which the boring apparatus of the
bathes these organs. This fluid embryo can be distinguished. (From R.
evaporates when desiccation Greeff.)
takes place, and is soon replaced
after rain; it forms no coagulum
when reagents are added to it, and it probably differs but little from
water. Floating in it are numerous corpuscles, whose number
increases with age. In well-fed Tardigrades the corpuscles are packed
with food-reserves, often of the same colour—green or brown—as the
 contents of the stomach, which
soon disappear when the little
creatures are starved.
The alimentary canal begins
with an oral cavity, which is in
many species surrounded by
chitinous rings. The number of
Fig. 251.—Echiniscus spinulosus, C.
these rings and their general
Sch., × about 200, seen from the side. arrangement are of systematic
(From Doyère.) importance. The oral cavity opens
behind into a fine tube lined with
chitin, very characteristic of the
Tardigrada, which has been termed the mouth-tube. By its side,
converging anteriorly, lie the two chitinous teeth, which may open
ventrally into the mouth-tube, as in Macrobiotus hufelandi and
Doyeria simplex, or may open directly into the oral cavity, as in
Echiniscus, Milnesium, and some species of Macrobiotus. In some of
the last named the tips of the teeth are hardened by a calcareous
deposit. The hinder end of each stylet or tooth is supported by a
second chitinous tooth-bearer,[375] and the movement of each is
controlled by three muscles, one of which, running forwards to the
mouth, helps to protrude the tooth, whilst the other two running
upwards and downwards to the sheath of the pharynx, direct in what
plane the tooth shall be moved.
The mouth-tube passes suddenly into the muscular sucking
pharynx, which is pierced by a continuation of its chitinous tube.
Roughly speaking, the pharynx is spherical; the great thickness of its
walls is due to radially arranged muscles which run from the
chitinous tube to a surrounding membrane. When the muscles
contract, the lumen of the tube is enlarged, and food, for the most
part liquid, is sucked in. Two large glands, composed of cells with
conspicuous nuclei, but with ill-defined cell outlines, pour their
contents into the mouth in close proximity to the exit of the teeth.
The secretion of the glands—often termed salivary glands—is said in
many cases to be poisonous.
The pharynx may be followed by a distinct oesophagus, or it may
pass almost immediately into the stomach, which consists of a layer
of six-sided cells arranged in very definite rows. In fully-fed
specimens these cells project into
the lumen with a well-rounded
contour. Posteriorly the stomach
contracts and passes into the
narrow rectum, which receives
anteriorly the products of the
excretory canals and the
reproductive organs, and thus
forms a cloaca. Its transversely
placed orifice lies between the last
pair of legs. The food of
Tardigrades is mainly the sap of
mosses and other humble plants,
the cell-walls of which are pierced
by the teeth of the little creatures.
The organs to which an
excretory function has been
attributed are a pair of lateral
caeca, which vary much in size
according as the possessor is well
or ill nourished. They recall the
Malpighian tubules of such Mites
as Tyroglyphus. Nothing
comparable in structure to
nephridia or to coxal glands has
been found. Fig. 252.—Macrobiotus schultzei, Gr.,
The muscles show a beautiful × 150. (Modified from Greeff.) a, The
symmetry. There are ventral, six inner papillae of the mouth; b, the
dorsal, and lateral bundles, and chitin-lined oesophagus; c, calcareous
others that move the limbs and spicule; d, muscle which moves the
spicule; e, muscular pharynx with
teeth, but the reader must be masticating plates; f, salivary glands; g,
referred to the works of Basse, stomach; h, ovary; i, median dorsal
Doyère,[376] and Plate[377] for the accessory gland; k, diverticula of
details of their arrangement. The rectum.
muscle-fibres are smooth.
The nervous system consists of a brain or supra-oesophageal
ganglion, whose structure was first elucidated by Plate, and a ventral
chain of four ganglia. Anteriorly the brain is rounded, and gives off a
 nerve to the skin; posteriorly each
half divides into two lobes, an
inner and an outer. The latter
bears the eye-spot when this is
present. Just below this eye a
slender nerve passes straight to
the first ventral ganglion. The
brain is continued round the oral
cavity as a thick nerve-ring, the
ventral part of which forms the
sub-oesophageal ganglion, united
by two longitudinal commissures
Fig. 253.—Brain of Macrobiotus to the first ventral ganglion. Thus
hufelandi, C. Sch., × about 350. (From the brain has two channels of
Plate.) Seen from the side. ap, Lobe of communication between it and
brain bearing the eye; ce, supra- the ventral nerve-cord on each
oesophageal ganglion; d, tooth; Ga,
first ventral ganglion; ga’, sub-
side, one by means of the slender
oesophageal ganglion; k, thickening of nerve above mentioned, and one
the epidermis round the mouth; oc, through the sub-oesophageal
eye-spot; oe, oesophagus; op, nerve ganglion. The ventral chain is
running from the ocular lobe of the composed of four ganglia
brain to the first ventral ganglion; ph, connected together by widely
pharynx. divaricated commissures. Each
ganglion gives off three pairs of
nerves, two to the ventral musculature, and one to the dorsal. The
terminations of these nerves in the muscles are very clearly seen in
these transparent little creatures, though there is still much dispute
as to their exact nature.
The older writers considered the Tardigrada as hermaphrodites,
but Plate and others have conclusively shown that they are bisexual,
at any rate in the genus Macrobiotus. The males are, however, much
rarer than the females. The reproductive organs of both sexes are
alike. Both ovary and testis are unpaired structures opening into the
intestine, and each is provided with a dorsal accessory gland placed
near its orifice. In the ovary many of the eggs are not destined to be
fertilised, but serve as nourishment for the more successful ova
which survive.
 No special circulatory or respiratory organs exist, and, as in many
other simple organisms, there is no connective tissue.
The segmentation of the egg in
M. macronyx is total and equal,
according to the observations of
von Erlanger.[378] A blastula,
followed by a gastrula, is formed.
The blastopore closes, but later
the anus appears at the same
spot. There are four pairs of
mesodermic diverticula which
give rise to the coelom and the
chief muscles. The reproductive
organs arise as an unpaired
diverticulum of the alimentary
canal, which also gives origin to
the Malpighian tubules. The
development is thus very
primitive and simple, and affords
no evidence of degeneration. Fig. 254.—Male reproductive organs of
With regard to their position in Macrobiotus hufelandi, C. Sch., ×
about 350. (From Plate.) a.ep,
the animal kingdom, writers on Epidermal thickening round anus; cl,
the Tardigrada are by no means cloaca; gl.d, accessory gland; gl.l,
agreed. O. F. Müller placed them Malpighian gland; st, stomach; te,
with the Mites; Schultze and testis; x, mother-cells of spermatozoa.
Ehrenberg near the Crustacea;
Dujardin and Doyère with the
Rotifers near the Annelids; and von Graff with the Myzostomidae
and the Pentastomida. Plate regards them as the lowest of all air-
breathing Arthropods, but he carefully guards himself against the
view that they retain the structure of the original Tracheates from
which later forms have been derived. He looks upon Tardigrades as a
side twig of the great Tracheate branch, but a twig which arises
nearer the base of the branch than any other existing forms. These
animals seem certainly to belong to the Arthropod phylum,
inasmuch as they are segmented, have feet ending in claws,
Malpighian tubules, and an entire absence of cilia. The second and
third of these features indicate a relationship with the Tracheate
groups; on the other hand there is an absence of paired sensory
appendages, and of mouth-parts. Von Erlanger has pointed out that
the Malpighian tubules, arising as they do from the mid-gut, are not
homologous with the Malpighian tubules of most Tracheates, and he
is inclined to place this group at the base or near the base of the
whole Arthropod phylum. They, however, show little resemblance to
any of the more primitive Crustacea. The matter must remain to a
large extent a matter of opinion, but there can be no doubt that the
Tardigrades show more marked affinities to the Arthropods than to
any other group of the animal kingdom.
Biology.—Spallanzani, who published in the year 1776 his
Opuscules de physique animale et végétale, was the first
satisfactorily to describe the phenomena of the desiccation of
Tardigrades, though the subject of the desiccation of Rotifers,
Nematodes, and Infusoria had attracted much notice, since
Leeuwenhoek had first drawn attention to it at the very beginning of
the century. In its natural state and in a damp atmosphere
Tardigrades live and move and have their being like other animals,
but if the surroundings dry up, or if one be isolated on a microscopic
slide and slowly allowed to dry, its movements cease, its body
shrinks, its skin becomes wrinkled, and at length it takes on the
appearance of a much weathered grain of sand in which no parts are
distinguishable. In this state, in which it may remain for years, its
only vital action must be respiration, and this must be reduced to a
minimum. When water is added it slowly revives, the body swells,
fills out, the legs project, and gradually it assumes its former plump
appearance. For a time it remains still, and is then in a very
favourable condition for observation, but soon it begins to move and
resumes its ordinary life which has been so curiously interrupted.
All Tardigrades have not this peculiar power of revivification—
anabiosis, Preyer calls it—it is confined to those species which live
amongst moss, and the process of desiccation must be slow and,
according to Lance,[379] the animal must be protected as much as
possible from direct contact with the air.
According to Plate, the Tardigrada are free from parasitic Metazoa,
which indeed could hardly find room in their minute bodies. They
are, however, freely attacked by Bacteria and other lowly vegetable
organisms, and these seem to flourish in the blood without
apparently producing any deleterious effects on the host. Plate also
records the occurrence of certain enigmatical spherical bodies which
were found in the blood or more usually in the cells of the stomach.
These bodies generally appeared when the Tardigrades were kept in
the same unchanged water for some weeks. Nothing certain is known
as to their nature or origin.
Systematic.—A good deal of work has recently been done by Mr.
James Murray on the Polar Tardigrades and on the Tardigrades of
Scotland, many of which have been collected by the staff of the Lake
Survey.[380] Over forty species have been described from North
Britain.
The following table of Classification is based on that drawn up by
Plate:—
 Table of Genera.
I. The claws of the legs are simple, without a second hook. If there
are several on the same foot they are alike in structure and size.
A. The legs are short and broad, each with at least two claws.

2–4 claws Gen. 1. ECHINISCUS, C. Sch. (Fig. 249).

7–9 claws Sub-gen. 1a. ECHINISCOIDES, Plate.

B. The legs are long and slender; each bears only one small
claw.

Gen. 2. LYDELLA, Doy.

II. The claws of the legs are all or partly two- or three-hooked.
Frequently they are of different lengths.
A. There are no processes or palps around the mouth.
I. The muscular sucking pharynx follows closely on the
mouth-tube.
α. The oral armature consists on each side of a stout
tooth and a transversely placed support.

Gen. 3. MACROBIOTUS, C. Sch.

(Fig. 252).

β. The oral armature consists on each side of a stylet-

like tooth without support.

Gen. 4. DOYERIA, Plate.

II. The mouth-tube is separated from the muscular

sucking pharynx by a short oesophagus.

Gen. 5. DIPHASCON, Plate (Fig.

255).
 B. Six short processes or palps surround the mouth, and
two others are placed a little farther back.

Gen. 6. MILNESIUM, Doy.

1. Genus ECHINISCUS (= EURYDIUM, Doy.).—The dorsal cuticle

is thick, and divided into a varying number of shields, which bear
thread- or spike-like projections. The anterior end forms a proboscis-
like extension of the body. Two red eye-spots. There are many
species, and the number has increased so rapidly in the last few years
that specialists are talking of splitting up the genus. E. arctomys,
Ehrb.; E. mutabilis, Murray; E. islandicus, Richters; E. gladiator,
Murray; E. wendti, Richters; E. reticulatus, Murray; E. oihonnae,
Richters; E. granulatus, Doy.; E. spitzbergensis, Scourfield;[381] E.
quadrispinosus, Richters; and E. muscicola, Plate, are all British.
More than one-half of these species are also Arctic, and E. arctomys
is in addition Antarctic. In fact, the group is a very cosmopolitan one.
The genus is also widely distributed vertically, specimens being
found in cities on the sea level and on mountains up to a height of
over 11,000 feet.
1a. Sub-genus ECHINISCOIDES differs from the preceding in the
number of the claws, the want of definition in the dorsal plates, and
in being marine. The single species E. sigismundi, M. Sch., is found
amongst algae in the North Sea (Ostend and Heligoland).
2. Genus LYDELLA.[382]—The long, thin legs of this genus have two
segments, and in other respects approach the Arthropod limb.
Marine. Plate suggests the name L. dujardini for the single species
known.
3. Genus MACROBIOTUS has a pigmented epidermis, but eye-
spots may be present or absent. The eggs are laid one at a time, or
many leave the body at once. They are either quite free or enclosed in
a cast-off cuticle. The genus is divided into many species and shows
signs of disruption. They mostly live amongst moss; but M.
macronyx, Doy., is said to live in fresh water. The following species
are recorded from North Britain: M. oberhäuseri, Doy.; M.
hufelandi, Schultze; M. zetlandicus, Murray; M. intermedius, Plate;
M. angusti, Murray; M. annulatus, Murray; M. tuberculatus, Plate;
M. sattleri, Richters; M. papillifer, Murray; M. coronifer, Richters;
 M. crenulatus, Richters; M.
harmsworthi, Murray; M.
orcadensis, Murray; M.
islandicus, Richters; M. dispar,
Murray; M. ambiguus, Murray;
M. pullari, Murray; M. hastatus,
Murray; M. dubius, Murray; M.
echinogenitus, Richters; M.
ornatus, Richters; M. macronyx?,
Doy.
4. Genus DOYERIA.—The teeth
of this genus have no support,
and the large salivary glands of
the foregoing genus are absent; in
other respects Doyeria, with the
single species Doyeria simplex,
Plate, resembles Macrobiotus,
and is usually to be found in
consort with M. hufelandi, C. Sch.
5. Genus DIPHASCON
resembles M. oberhäuseri, Doy.,
but an oesophagus separates the
mouth-tube from the sucking
pharynx, and the oral armature is
weak. The following species are
British, the first named being
very cosmopolitan, being found at
both Poles, in Chili, Europe, and
Asia: D. chilenense, Plate; D.
Fig. 255.—Diphascon chilenense, Plate, scoticum, Murray; D. bullatum,
× about 100. (From Plate.) ce, Brain; k, Murray; D. angustatum, Murray;
thickening of the epidermis above the
mouth; o, egg; oe, oesophagus; p,?
D. oculatum, Murray; D.
salivary glands; ph, pharynx; sa, blood alpinum, Murray; D.
corpuscles; st, stomach. spitzbergense, Murray.
6. Genus MILNESIUM has a
soft oral armature, and the teeth
open straight into the mouth. A lens can usually be distinguished in
the eyes. Two species have been described, M. tardigradum, Doy.,
British, and M. alpigenum, Ehrb. Bruce and Richters consider that
these two species are identical.
 CHAPTER XX
[383]
PENTASTOMIDA

OCCURRENCE—ECONOMIC IMPORTANCE—STRUCTURE
—DEVELOPMENT AND LIFE-HISTORY—SYSTEMATIC

Pentastomids are unpleasant-looking, fluke-like or worm-like

animals, which pass their adult lives in the nasal cavities, frontal
sinuses, and lungs of flesh-eating animals, such as the Carnivora,
Crocodiles, and Snakes; more rarely in Lizards, Birds, or Fishes.
From these retreats their eggs or larvae are sneezed out or coughed
up, or in some other way expelled from the body of their primary
host, and then if they are eaten, as they may well be if they fall on
grass, by some vegetable-feeding or omnivorous animal, they
undergo a further development. If uneaten the eggs die. When once
in the stomach of the second host, the egg-shell is dissolved and a
larva emerges (Fig. 260, p. 494), which bores through the stomach-
wall and comes to rest in a cyst in some of the neighbouring viscera.
Here, with occasional wanderings which may prove fatal to the host,
it matures, and should the second host be eaten by one of the first,
the encysted form escapes, makes its way to the nasal chambers or
lungs, and attaching itself by means of its two pairs of hooks, comes
to rest on some surface capable of affording nutriment. Having once
taken up its position the female seldom moves, but the males, which
are smaller than the females, are more active. They move about in
search of a mate. Further, should the host die, both sexes, after the
manner of parasites, attempt to leave the body. Like most animals
who live entirely in the dark they develop no pigment, and have a
whitish, blanched appearance.
The only species of Pentastomid which has any economic
importance is Linguatula taenioides of Lamarck, which is found in
the nose of the dog, and much more rarely in the same position in
the horse, mule, goat, sheep, and man. It is a comparatively rare
parasite, but occurred in about 10 per cent of the 630 dogs in which
it was sought at the laboratory of Alfort, near Paris, and in 5 out of
60 dogs examined at Toulouse. The symptoms caused by the
presence of these parasites are not usually very severe, though cases
have been recorded where they have caused asphyxia. The larval
stages occur in the rabbit, sheep, ox, deer, guinea-pig, hare, rat,
horse, camel, and man, and by their wandering through the tissues
may set up peritonitis and other troubles.
As in the Cestoda, which they so closely resemble in their life-
history, the nomenclature of the Pentastomids has been complicated
by their double life. For long the larval form of L. taenioides was
known by different names in different hosts, e.g. Pentastoma
denticulatum, Rud., when found in the goat, P. serratum, Fröhlich,
when found in the hare, P. emarginatum when found in the guinea-
pig, and so on. In the systematic section of this article some of the
species mentioned are known in the adult state, some in the larval,
and in only a few has the life-history been fully worked out.
Structure.[384]—The body of a Pentastomid is usually white,
though in the living condition it may be tinged red by the colour of
the blood upon which it lives. The anterior end, which bears the
mouth and the hooks (Fig. 256), has no rings; this has been termed
the cephalothorax. The rest of the body, sometimes called the
abdomen, is ringed, and each annulus is divided into an anterior half
dotted with the pores of certain epidermal glands and a hinder part
of the ring in which these are absent.
On the ventral surface of the cephalothorax, in the middle line, lies
the mouth, elevated on an oral papilla, and on each side of the mouth
are a pair of hooks whose bases are sunk in pits. The hooks can be
protruded from the pits, and serve as organs of attachment. Their
shape has some systematic value.
There are a pair of peculiar papillae which bear the openings of the
“hook-glands,” lying just in front of the pairs of hooks, and other
smaller papillae are arranged in pairs on the cephalothorax and
anterior annuli. The entire body is covered by a cuticle which is
tucked in at the several orifices. This is secreted by a continuous
layer of ectoderm cells. Some of these subcuticular cells are
aggregated together to form very definite glands opening through the
cuticle by pores which have somewhat unfortunately received the
name of stigmata. Spencer attributes to these glands a general
excretory function. There is, however, a very special pair of glands,
the hook-glands, which extend almost from one end to the other of
the body; anteriorly these two lateral glands unite and form the
head-gland (Fig. 257). From this
on each side three ducts pass, one
of which opens to the surface on
the primary papilla; the other two
ducts open at the base of the two
hooks which lie on each side of
the mouth. Leuckart has
suggested that these important
glands secrete some fluid like the
irritating saliva of a Mosquito
which induces an increased flow
of blood to the place where it is of
use to the parasite. Spencer,
however, regards the secretion as
having, like the secretion of the
so-called salivary cells of the
Leech, a retarding action on the
coagulation of the blood of the
host.
The muscles of Pentastomids
are striated. There is a circular
layer within the subcuticular
cells, and within this a
longitudinal layer and an oblique
layer which runs across the body-
cavity from the dorso-lateral
surface to the mid-ventral line, a Fig. 256.—Porocephalus annulatus,
primitive arrangement which Baird. A, Ventral view of head, × 6; B,
recalls the similar division of the ventral view of animal, × 2.
body-cavity into three chambers
in Peripatus and in many Chaetopods. Besides these there are
certain muscles which move the hooks and other structures.
The mouth opens into a pharynx which runs upwards and then
backwards to open into the oesophagus (Fig. 257). Certain muscles
attached to these parts enlarge their cavities, and thus give rise to a
sucking action by whose force the blood of the host is taken into the
alimentary canal. The oesophagus opens by a funnel-shaped valve
into the capacious stomach or mid-gut, which stretches through the
body to end in a short rectum or hind-gut. The anus is terminal.

Fig. 257.—Diagrammatic representation of the alimentary,

secretory, nervous, and reproductive systems of a male
Porocephalus teretiusculus, seen from the side. The nerves are
represented by solid black lines. (From W. Baldwin Spencer.)

1, Head-gland; 2, testis; 3, hook-gland; 4, hind-gut; 5, mid-gut; 6,

ejaculatory
duct; 7, vesicula seminalis; 8, vas deferens; 9, dilator-rod sac; 10,
cirrus-bulb;
11, cirrus-sac; 12, fore-gut; 13, oral papillae.

There appears to be no trace of circulatory or respiratory organs,

whilst the function usually exercised by the nephridia or Malpighian
tubules or by coxal glands, of removing waste nitrogenous matter,
seems, according to Spencer, to be transferred to the skin-glands.
The nervous system is aggregated into a large ventral ganglion
which lies behind the oesophagus. It gives off a narrow band devoid
of ganglion-cells, which encircles that tube. It also gives off eight
nerves supplying various parts, and is continued backward as a ninth
pair of prolongations which, running along the ventral surface, reach
almost to the end of the body (Fig. 257). The only sense-organs
known are certain paired papillae on the head, which is the portion
that most closely comes in contact with the tissues of the host.
Pentastomids are bisexual. The males are as a rule much less
numerous and considerably smaller than the females, although the
number of annuli may be greater.
The ovary consists of a single tube closed behind. This is supported
by a median mesentery. Anteriorly the ovary passes into a right and
left oviduct, which, traversing the large hook-gland, encircle the
alimentary canal and the two posterior nerves (Fig. 258). They then
unite, and at their point of union they receive the ducts of the two
spermathecae, usually found packed with spermatozoa. Having
received the orifices of the spermatheca, the united oviducts are
continued backward as the uterus, a highly-coiled tube in which the
fertilised eggs are stored. These are very numerous; Leuckart
estimated that a single female may contain half a million eggs. The
uterus opens to the exterior in the mid-ventral line a short distance—
in P. teretiusculus on the last ring but seven—in front of the terminal
anus. In L. taenioides the eggs begin to be laid in the mucus of the
nose some six months after the parasite has taken up its position.

Fig. 258.—Diagrammatic representation of the alimentary,

secretory, nervous, and reproductive systems of a female
Porocephalus teretiusculus, seen from the side. The nerves are
represented by solid black lines. (From W. Baldwin Spencer.)

1, Head-gland; 2, oviduct; 3, hook-gland; 4, mid-gut; 5, ovary; 6,

hind-gut; 7,
vagina; 8, uterus; 9, accessory gland; 10, spermatheca.

The testis is a single tube occupying in the male a position similar

to that of the ovary in the female. Anteriorly it opens into two
vesiculae seminales, which, like the oviducts, pierce the hook-glands
and encircle the alimentary canal (Fig. 257). Each vesicula passes
into a vas deferens with a cuticular lining. Each vas deferens also
receives the orifice of a muscular caecal ejaculatory duct, which,
crowded with mature spermatozoa, stretches back through the body.
Anteriorly the vas deferens passes into a cirrus-bulb, which is joined
by a cirrus-sac on one side and a dilator-rod sac on the other,
structures containing organs that assist in introducing the
spermatozoa into the female. The two tubes then unite, and having