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Conservation Biology
in Sub-Saharan Africa
Copyright © 2019. Open Book Publishers. All rights reserved.
IN SUB-SAHARAN AFRICA
CONSERVATION BIOLOGY
Copyright © 2019. Open Book Publishers. All rights reserved.
Conservation Biology
in Sub-Saharan Africa
This work is licensed under a Creative Commons Attribution 4.0 International license (CC BY 4.0). This
license allows you to share, copy, distribute and transmit the text; to adapt the text and to make commercial
use of the text providing attribution is made to the authors (but not in any way that suggests that they
endorse you or your use of the work). Attribution should include the following information:
John W. Wilson and Richard B. Primack, Conservation Biology in Sub-Saharan Africa. Cambridge, UK: Open
Book Publishers, 2019, https://doi.org/10.11647/OBP.0177
In order to access detailed and updated information on the license, please visit, https://doi.org/10.11647/
OBP.0177#copyright
All external links were active at the time of publication unless otherwise stated and have been archived via
the Internet Archive Wayback Machine at https://archive.org/web
Updated digital material and resources associated with this volume are available at https://doi.org/10.11647/
OBP.0177#resources
Licencing for images obtained online, many of which underwent minor stylistic adjustments, were correct
at the time of publication.
Every effort has been made to identify and contact copyright holders and any omission or error will be
corrected if notification is made to the publisher.
The Lounsbery Foundation has generously contributed to the publication of this volume.
3. What is Biodiversity?
3.1 Species Diversity 62
3.1.1 What is a species? 64
3.2 Genetic Diversity 65
3.3 Ecosystem Diversity 69
3.4 Patterns of Biodiversity 71
3.4.1 Challenging species identifications 72
3.4.2 Implications of challenging species identifications 73
3.4.3 Measuring species diversity 79
3.4.4 How many species exist? 81
Copyright © 2019. Open Book Publishers. All rights reserved.
Bibliography 126
8. Extinction Is Forever
8.1 What is Extinction? 259
8.2 Rates of Extinction 259
8.3 When is a Species Extinct? 260
8.4 History of Extinctions in Sub-Saharan Africa 263
8.5 Which Species are at Risk of Extinction? 271
8.5.1 Course-filter assessments 274
8.6 Characteristics of Threatened Species 274
8.7 Problems of Small Populations 277
8.7.1 Loss of genetic diversity 277
Genetic drift 277
Inbreeding depression 278
Outbreeding depression 280
Population bottlenecks 281
8.7.2 Demographic stochasticity 281
8.7.3 Environmental stochasticity and catastrophes 282
8.7.4 The extinction vortex 284
8.7.5 Is there any hope for small populations? 285
8.8 Is De-extinction a Solution? 285
8.9 Summary 289
8.10 Topics for Discussion 289
8.11 Suggested Readings 290
Bibliography 291
14.1.2 The impact of logging, mining, and other extractive industries 530
14.2 Smart Development Outside Conservation Areas 532
14.3 Linking Conservation to Socio-Economic Development 534
14.4 Confronting Human-Wildlife Conflict 539
14.4.1 Dealing with predators 540
14.4.2 Dealing with crop raiders 541
14.4.3 Concluding thoughts on human-wildlife conflict 542
14.5 Summary 542
Contents xvii
Appendix A 589
Selected Sources of Information
Appendix B 593
Selected Environmental Organisations
Appendix C 607
Obtaining Conservation Funding
Copyright © 2019. Open Book Publishers. All rights reserved.
Appendix D 613
Environmental Calendar
Glossary 615
Index 645
Copyright © 2019. Open Book Publishers. All rights reserved.
List of Boxes
Box 1.2 The Okapi Wildlife Reserve: Protecting Nature and Providing for People 14
Rosmarie Ruf & Marcel Enckoto
Box 2.4 Malawi: No Longer a Weak Link in the Elephant Ivory Trafficking Chain? 48
Jonathan Vaughan
Box 4.4 Conservation Lessons from the Asian and African Vulture Crises 110
Ara Monadjem
Box 5.1 The Importance of Liberia’s Forest Network to the Survival of 136
the Pygmy Hippopotamus
Mary Molokwu-Odozi and Kathryn Phillips
Box 5.2 The Conservation and Exploitation of East African Plants 146
John R. S. Tabuti
Box 5.3 Migratory Birds of Africa: The Largest of the Last Great Migrations? 150
Abraham J. Miller-Rushing and John W. Wilson
Box 5.4 Saving Critically Endangered Ground Nesting Birds from Habitat Loss 153
Bruktawit Abdu Mahamued
Box 6.1 Does Oil Palm Agriculture Threaten Biological Diversity 170
in Equatorial Africa?
Abraham J. Miller-Rushing
Box 6.3 Habitat Alteration, Climate Change, and Mosquito-Borne Diseases 190
Kevin Njabo
Box 7.1 Solving Seabird Bycatch Problems: From Theory to Practice 221
Copyright © 2019. Open Book Publishers. All rights reserved.
Ross Wanless
Box 8.1 Pleistocene Extinctions: Climate Change, Hominin Predation, or Both? 263
David H.M. Cumming
Box 8.3 Fenced Reserves Conserving Cheetahs and African Wild Dogs 286
in South Africa
Kelly Marnewick
Box 9.1 The Role of Biodiversity Inventories in the Management of Gorongosa 300
National Park
Marc Stalmans and Piotr Naskrecki
Box 9.2 Sea Turtle Conservation along Africa’s Atlantic Coast 303
Angela Formia
Box 9.3 Sustainably Harvesting Fruit Bats Through Better Understanding 315
of Life Histories
David T. S. Hayman
Box 10.2 Remote Sensing and Spatial Analysis for African Conservation 335
Barend F. N. Erasmus
Box 11.1 The Overlooked Role of Behavioural Ecology in the Conservation 378
of African Mammals
Adrian M Shrader
Box 11.4 Saving the Northern White Rhinoceros with Assisted Reproduction 414
Technologies
Morné de la Rey
Box 12.1 Insect Biodiversity Helps Solve African Wildlife Crimes 442
Martin H. Villet
Box 13.1 Mpala Research Centre: A Living Laboratory for (More than Just) 467
Scientists
Anchal Padukone and Dino J. Martins
Box 13.2 Identifying Key Sites for Conservation in the Albertine Rift 475
Andrew J. Plumptre
Box 13.3 Marine Protected Areas in East Africa and the Western Indian Ocean 481
Abraham J. Miller-Rushing
Box 13.4 Zoning: Something for Everyone in the Forests of Dzangha-Sangha 487
Richard Carroll
Box 14.1 Traditional People and Conservation: Turning the Page 517
Abraham J. Miller-Rushing and John W. Wilson
Copyright © 2019. Open Book Publishers. All rights reserved.
Box 14.2 Importance of Protected Areas in Cities: Insights from the City 523
of Cape Town
Pippin M. L. Anderson
Box 15.3 Tracking Species in Space and Time: Citizen Science in Africa 570
Phoebe Barnard
Even for those of us who are one step removed from nature in our present-day lives,
looking back on places we called home as children can reveal disturbing truths. This is
certainly true for me. I grew up in a small mining town in Zimbabwe called Kadoma,
and as soon as school was out, my parents would bundle me off to Mhondoro, a rural
area to the east.
I loved it there and happily spent hours herding cattle across the quintessential
untamed African savannah. I have many fond memories of swimming and catching
fish in Mhondoro too. I recall how the surrounding forests were lush, teeming with
wildlife; rivers were bountiful, full of fish and fowl in the pristine fresh water. City
folk went home with their hands full of gifts from nature and agricultural fields.
Today, when I return to my village in Mhondoro, my heart breaks. The lush forests
and wildlife are gone, replaced with barren fields and a whimpering stream where the
river once ran. I now bring my own food and bottled water when I visit. And worst of
all, the people of Mhondoro who I had always associated with nature’s abundance are
today poor and disenfranchised and have few if any options for bettering their lives.
Tragically, the story of my village is shared by thousands of villages across
Africa that are suffering the worst impacts of climate change, population growth
and harmful development choices. Faced with the challenge of feeding their families
and generating cash incomes, farmers, like those of Mhondoro, end up expanding
their crops increasingly deeper into wild lands and forests. These encroachments not
only bring their families into dangerous conflict with wildlife, they simultaneously
endanger and destroy the forests and fertile soils that would otherwise support their
agricultural bounty.
Copyright © 2019. Open Book Publishers. All rights reserved.
new deal for nature, we must support research capacity and skills building within
its populations, including investments in faculty and research leaders, facilities and
infrastructure, and expanding career opportunities for budding researchers to apply
their findings in real world settings.
Fortunately, there is now widespread acknowledgement that African researchers
are best placed to ask questions and find solutions to the challenges facing Africa.
Hence, we are witnessing a slow move away from the notion that researchers from
high income countries must be parachuted in to identify and address the continent’s
problems.
This textbook, Conservation Biology in Sub-Saharan Africa, is a critical first step
as it goes a long way towards focusing attention to the urgent need to define the
nature of the problem and develop practical, context specific solutions to Africa’s
many environmental challenges. It discusses how our lives are inextricably linked
to a healthy environment, explores how inclusive conservation can provide greater
benefits to all, and illustrates how grassroots action can ensure that nature’s many
beneficial contributions will remain available for generations to come.
By being distributed for free, this textbook ensures that its readership can include
those who stand to benefit the greatest, including African researchers and practitioners.
Open access textbooks like it are critical to expanding access to African research and
improving intra-African research collaboration and capacity.
If we are to “leave no one behind,” as agreed by global leaders in 2015 and
encapsulated in Sustainable Development Goals, farmers like those from Mhondoro
will also need access to the best available information, science and solutions. That is a
deal that we owe them—and the many people who call Africa their home. This book
makes an important contribution to the challenge. I hope that, thanks to the efforts of
the experts featured in this textbook and others, one day when I return to Mhondoro,
it will more closely resemble its prior self that I loved as a child.
Maxwell Gomera
Director: Biodiversity and Ecosystem Services Branch
UN Environment
Copyright © 2019. Open Book Publishers. All rights reserved.
Preface
We are excited to present the very first conservation biology textbook dedicated
entirely to an African audience. The need for this work has never been more pressing
than now. Africa has some of the fastest growing human populations on Earth.
This growth, together with a much-needed push for development to ensure that all
Africans can live healthy and prosperous lives, exerts unsustainable pressure on the
region’s rich and unique biological treasures. Consequently, Africa is losing its natural
heritage faster than ever before. It is sobering to consider that there is a very real
risk that our children may never have the opportunity to see gorillas, rhinoceros, or
elephants in the wild.
To address this alarming loss of Africa’s natural heritage, there is an urgent need
to produce the next cohort of well-trained conservation leaders, able to confront
conservation challenges head-on, and to secure a sustainable future for all. This effort
starts early, by exposing children from a young age to the wonders of the natural
world. But it is also important to ensure that those children who later choose a career
in biodiversity conservation are well-prepared for the road ahead. To facilitate this
capacity building, we have compiled this textbook, designed for use in conservation
biology courses, and as a supplemental text for other courses in the natural sciences
and environmental policy. While the main target audience for this book is early-
career conservationists, we strived for a balance between theory, empirical data, and
practical guidelines to also make the book a valuable resource for mid- and late-career
professionals. To further remove obstacles to training, we made every effort to ensure
that this work is accessible to as wide an audience as possible. For that reason, we are
making this textbook available for free, under a Creative Commons (CC BY) license,
Copyright © 2019. Open Book Publishers. All rights reserved.
to guarantee the rights for anyone to use and spread this work to whoever wishes to
make a difference in the future of Africa’s biodiversity and its people.
Scope
This textbook focuses on the Afrotropics, one of Earth’s eight major terrestrial
ecozones. This area includes continental Africa south of the Sahara Desert, continental
islands (e.g. the Seychelles) that drifted away from Africa millions of years ago, and
oceanic islands with a volcanic origin (e.g. the Comoros archipelago, São Tomé, and
xxviii Conservation Biology in Sub-Saharan Africa
Príncipe) which share many biological characteristics with the Afrotropics. This area
is also generally known as Sub-Saharan Africa, which we use throughout this book
as a convenient and acceptable way to designate this ecologically (African parts of
Afrotropics) and geographically (Africa south of the Sahara) distinct region.
Deviating slightly from the typical scope of some other books focussed on the
Afrotropics, this textbook does not cover south-west Arabia—these areas, together
with North Africa, are covered in a sister textbook to this one, published in Arabic
(Primack and El-Demerdash, 2003). Similarly, this book excludes Madagascar and the
Mascarene Islands, which are covered in two sister textbooks published in French
(Primack and Ratsirarson, 2005; Primack et al., 2012). The text does cover a few areas
Copyright © 2019. Open Book Publishers. All rights reserved.
not usually considered part of the Afrotropics, but which share several affinities with
the Afrotropical region. These additional areas include oceans within 200 nautical
miles from Sub-Saharan Africa, and oceanic islands in the Atlantic that are usually
treated as part of the Palearctic realm, namely Cabo Verde, St. Helena, Ascension, and
the Tristan da Cunha archipelago.
summarises the threat status of many species on Earth. The classification system
used to compile this database is discussed in detail in Section 8.5. We foreshadow
this discussion by alerting readers that the threat status of each assessed species
that is mentioned in the text is indicated with one of the seven acronyms mentioned
below, right after its scientific name. To facilitate this indication, we generally follow
the IUCN’s taxonomy in this textbook. We are fully aware that the lag time between
taxonomic updates and IUCN assessment updates may create the appearance that this
textbook’s taxonomy might sometimes be outdated. We made every effort to highlight
important taxonomic discrepancies when relevant to the text. For common names, we
tried to use the most-widely used terms across Africa.
EX Extinct
EW Extinct in the Wild
EN Endangered
VU Vulnerable
NT Near Threatened
LC Least Concern
DD Data Deficient
of specialist terms listed in the glossary. Also included are over 50 Boxes, authored
by conservation researchers and practitioners from governments, universities, and
nongovernmental organisations in West, Central, East, and Southern Africa. These
case studies cover interdisciplinary topics such as public health, sacred spaces, energy,
agriculture, law, sustainable development, and leadership. They expose readers to
the voices of conservationists in the region, provide compelling examples of on-the-
ground work, and offer insights into real-life conservation issues readers may face in
their careers.
In each chapter, following the main body of text, there is a brief summary of main
take-home messages, a list of discussion questions, and a list of suggested readings.
xxx Conservation Biology in Sub-Saharan Africa
The discussion questions are formulated in such a way that there are no definitive right
or wrong answers—rather, they are meant to stimulate discussion among readers so
that they can develop their own conservation philosophies. We tried to restrict the
suggested reading lists to works that are freely available either on publisher websites
or online depositories accessible through Google Scholar (https://scholar.google.
com). Note that the suggested readings are not meant to be absolute; in fact, they
should be adapted to meet local contexts and syllabus requirements. Each chapter
concludes with an extensive bibliography, which serves to provide a starting point for
readers interested in specific conservation topics, and lecturers interested in adapting
the suggested reading list. The textbook concludes with four appendices, meant to
encourage readers to take the field’s activist spirit to heart.
While we have made every attempt to ensure that the content is current and
comprehensive, we recognise that mistakes do creep in; the field of conservation
biology is currently also rapidly evolving, including being more inclusive of previously
marginalised communities. We thus plan on continue updating this text, and welcome
comments and suggestions from readers who share our interest in protecting Sub-
Saharan Africa’s natural heritage.
Bibliography
Primack, R.B., and J. Ratsirarson. 2005. Principe de Base de la Conservation de la Biodiversité (in
French) (Antananarivo: University of Antananarivo).
Primack, R.B., F. Sarrazin, and J. Lecompte. 2012. Biologie de la Conservation (in French) (Paris:
Dunod).
Primack R.B., and M. El-Demerdash. 2003. Essentials of Conservation Biology (in Arabic) (Cairo:
Mars Publications).
Copyright © 2019. Open Book Publishers. All rights reserved.
Acknowledgements
We sincerely appreciate the contributions of everyone who helped ensure that this
project was a success. In particular, we would like to thank Peter Ryan for his guidance
and support throughout the project, Abe Miller-Rushing for writing several boxes
and providing editorial input, and the Richard Lounsbery Foundation for funding
support. Peter Ryan, David Cumming, and Adrian Shrader provided invaluable
reviews to ensure the content is accurate and pitched at the right level for conservation
students. We would also like to recognise the support from Nigel Bennett, Duan
Biggs, Meg Boeni, Israel Borokini, and Mike Scantlebury at various stages of the
project. Additional input on ex situ conservation was provided by Dalia Amor Conde,
Johanna Staerk, and Simeon Smeele at Species360, while Peter Maniloff provided
input on environmental economics. We thank all our photographers and box authors
for their willingness to contribute to this work. Alessandra Tosi, Rupert Gatti, and the
rest of the staff at Open Book Publishers helped to transform the manuscript into a
finished product. Lastly, a special thanks to Lesley Starke and Margaret Primack for
their encouragement, support, advice, and input throughout the project.
Copyright © 2019. Open Book Publishers. All rights reserved.
Copyright © 2019. Open Book Publishers. All rights reserved.
List of Acronyms
East Africa’s great migration is one of the most famous wildlife spectacles on Earth. Each year, tens of thousands of
tourists from around the world flock to the region to see the 1.7 million common wildebeest (Connochaetes taurinus, LC)
and hundreds of thousands of other plains mammals make their way from Tanzania’s Ngorongoro Conservation Area,
through the Serengeti Plains, to Kenya’s Maasai Mara National Reserve. Photograph by Daniel Rosengren, https://
commons.wikimedia.org/wiki/File:Wildebeest_Migration_in_Serengeti_National_Park,_Tanzania.jpg, CC BY 4.0.
To survive and prosper, people use natural resources. They harvest and use oil, water,
and wildlife products, and convert natural ecosystems for agriculture, cities, roads,
and industrial activities. This consumption, which reduces natural habitat and the
associated wildlife populations, is intensifying because of the demands of a rapidly
increasing human population. Consumption of resources also increases as countries
develop and industrialise: the average citizen of the USA uses five times more
Chapter 1 | What is Conservation Biology? 3
The first two goals describe typical scientific research investigating objective facts.
The third goal, however, is a part of what makes conservation biology a normative
discipline; that is, conservation biology incorporates human values, not just facts,
4 Conservation Biology in Sub-Saharan Africa
to understand and achieve its value-laden goals (Lindenmayer and Hunter, 2010).
In this sense, conservation biology is related to environmentalism, in which people
aim to protect the natural environment for its own sake (see Section 4.3.2). However,
conservation biology is at its core a scientific discipline; it is founded on scientific
principles. This is not to say you must be a scientist to practice conservation biology;
there are many people who are not scientists who apply the principles of conservation
biology in their professional and personal lives.
The emergence of conservation biology as a distinct scientific field in the 1970s has
given rise to the formation of various formal societies representing the field in a united
voice. Most notable among these is the Society for Conservation Biology (SCB, Figure
1.2), which is a non-profit international professional organisation with a mission to
advance “the science and practice of conserving the Earth’s biological diversity”. To
facilitate opportunities where like-minded people can share ideas locally, the SCB
has regional branches, including an active Africa Section (http://conbio.org/groups/
sections/africa) which hosts regular conferences. In addition to the SCB, a great
number of other local, national, and regional conservation organisations also exist
and act as mouthpieces for grassroots movements and as custodians of nature. Many
of these groups focus on specific animals or local protected areas. Others organically
adapt their missions and visions in response to a specific need or threat. For example,
established in 1913 as an exchange forum between collectors of rare plants, the
Botanical Society of South Africa now actively works toward protecting those rare
plants in their natural habitats.
Conservation biology also has a history of adapting to new challenges. The very first
conservation activities, in Africa and beyond, were geared towards securing the rights
to valuable natural resources for people in powerful positions, such as kings and tribal
chiefs, enforced through a strictly adherence to cultural norms and customary laws
(Section 2.2). But as a growing human population expanded its influence on the
environment, and wildlife started to decline, earliest conservation models gradually
shifted towards fortress conservation approaches (Wilshusen et al., 2002) which
aimed to shield wildlife from people by setting aside protected areas where human
activities were strictly controlled.
Chapter 1 | What is Conservation Biology? 5
Figure 1.A A park ranger from the Uganda Wildlife Authority teaching HUGO community vol-
unteers how to collect faecal samples from gorilla night nests during a CTPH training workshop.
Photograph by CTPH, CC BY 4.0.
Our experience in initiating and managing PHE programmes for the past
10 years has taught us several lessons. One of the most important lessons to
ensure project sustainability is to regularly engage with local leaders and the
government. The Uganda Wildlife Authority, Uganda’s Ministry of Health, and
local health centres all attend CTPH meetings with VHCTs. Attendance by and
representation of these groups not only informs them of our activities, but also
Chapter 1 | What is Conservation Biology? 7
provides a platform to inform or train the VHCTs in what they would like them
to disseminate to the local communities.
We have also learnt that PHE-implementing partners and projects need to
be well-suited to each other and each site; this remains true even though health
needs are often the same, regardless of the location. For example, at Mount Elgon
National Park in Uganda, we found that training VHCTs in reducing conflict with
park management played a key role in changing community attitudes toward
conservation. In contrast, at Virunga National Park, Democratic Republic of the
Congo (DRC), we found that VHCTs needed to work more closely with local
health centres to prevent disease transmission between people and gorillas, and
to promote family planning in a largely Catholic country.
Lastly, we found that establishing income-generating projects for groups
rather than individuals was key to sustaining VHCT networks and programme
goals beyond donor funding cycles where we have had no volunteer dropouts
in the first 10 years of initiating the PHE programme. These key components
were accomplished by initiating livestock group enterprises and by encouraging
VHCT volunteers to invest generated income into Village Saving and Loan
Associations (see http://www.care.org/vsla).
Yet, as we consider how to best invest limited conservation resources, some difficult
questions arise. With seemingly more work to be done than can be accomplished,
should we let some species go extinct (Bottrill et al., 2008)? Which species? Who decides?
How can we even dare to think that we can play god? Such questions predictably
bring about strongly opinionated and emotional debate (Soulé, 2013 vs. Marvier, 2014;
Tallis and Lubchenco, 2014). Given the successful track record of fortress conservation
initiatives in preventing extinctions despite limited budgets (Young et al., 2014), as
well as the promising progress of more complex people-centred initiatives (Pooley et
al., 2014), it seems clear that conservation relies on some balance between these two
conservation philosophies (Sodhi et al., 2011). Conservation biologists of tomorrow
will be able to fine-tune the balance between these strategies by closely inspecting the
successes and failures of our actions today.
Copyright © 2019. Open Book Publishers. All rights reserved.
can influence public opinion and policy. As a starting point, those interested
in this role can join a conservation organisation to learn more about broader
issues. They could also use their personal networks to form alliances with
lawyers, citizen groups, and politicians.
2012). That is, conservation biologists are often required to take creative steps to
respond to imminent threats, typically without a complete knowledge of the systems
requiring attention. Conservation scientists must also articulate long-term visions for
conservation beyond solving immediate problems.
To be effective, conservation biologists must demonstrate the relevance of their
findings to a range of stakeholders. To be successful in this task, the importance of
sound scientific principles cannot be over-emphasised. Nature is a complex network
of many interdependent connections and feedback loops. Science is underpinned by
principles that provide conservationists the necessary quantitative and qualitative
tools to better measure and control for all these different aspects of biodiversity. Such
10 Conservation Biology in Sub-Saharan Africa
Figure 1.3 Conservation biology draws from many other sciences to protect biodiversity. It is closely related
to natural resource management, which aims to manage biodiversity primarily for the benefit of humans.
Integrated conservation and development projects (ICDP) are projects that manage nature for the benefit of
both humans and biodiversity. After Kareiva and Marvier, 2012; Temple, 1991, CC BY 4.0.