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Conservation Biology
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viii | Contents
4 Global Patterns and Drivers of Biodiversity

Biodiversity through Geologic Time
Formation of early life 82
82 Box 4.3 Case in Point
81
Discovery of the Human Biome 92
Radiation events 82 Realms, biomes, and ecoregions 93
Historical extinctions 84 The most diverse ecoregions 93
Global Patterns of Biodiversity 87 Biodiversity hotspots 96
How many species are there? 87 Latitudinal gradients 97
Drivers of Biodiversity 99
Box 4.1 Case in Point
Discovery of a New Way to Make Life 89 Evolutionary drivers 99
Ecological drivers 105
Box 4.2 Success Story
Synopsis 110
Rediscovery of an Extinct Species 90
Part II Importance of Biodiversity 115
5 The Many Values of Biodiversity

Values and Ethics 118
Types of Value Systems 119
117
Instrumental values 126
Relational values 129
Intrinsic values 120 Ethical Worldviews 135
Anthropocentrism 135
The Public Makeover of Sharks 124 Biocentrism 136
Ecocentrism 137
Religion and Conservation 126
6 Biodiversity and Ecosystem Services 141

What Are Ecosystem Services? 143
History of Ecosystem Services 143
Wetland Loss and Hurricane Katrina 163
Ecosystem valuation 144 Cultural services 164
Biodiversity and ecosystem function 146 Alternative frameworks 168
The Millennium Ecosystem Assessment 147 Biotic Control of Ecosystem Services 169
Types of Ecosystem Services 149 Functional traits 170
Supporting services 150 Biodiversity 171
Provisioning services 153 Box 6.3 Challenges & Opportunities
Regulating services 157 The Dilution Effect 175
Box 6.1 Success Story Ecosystem Markets and Payments for Ecosystem
How New York City Keeps Its Drinking Services 177
Water Clean 159
7 Ecological Economics 181

Principles of Ecological Economics 182
Decision-Making Analyses 185
Total Economic Value (TEV) Framework
Direct use value 193
192
Cost-effectiveness analysis 186 Indirect use value 194

Cost-benefit analysis 187 Option value 194
Multicriteria decision analysis 190 Existence value 194

Contents | ix
Box 7.1 Case in Point Surrogate market techniques 203

The Option Value of Biodiversity 195 Cost-based techniques 204
Estimates of TEV 196 Stated preference methods 206
Methods of Valuing Ecosystem Services 198 Box 7.3 Success Story
Revealed preference methods 198 Contingent Valuation Helps Reduce Human-Wildlife
Real market techniques 199 Conflict 208
Benefits transfer method 209
Box 7.2 Conservation in Practice
An Example of the Market Pricing Method: One Final Comment 209
Fisheries and Pollution 201
Box 7.4 Challenges & Opportunities
Taking Anthropocentrism to the Extreme 210
Part III Threats to Biodiversity 213
8 Extinction 215
Global Patterns of Endangerment
The IUCN Red List 216
216 Local Changes in Biodiversity
Species-area relationships 230
230
Box 8.1 Conservation in Practice Empirical measures 233

The IUCN Red List 217 Better monitoring and assessment programs 236
Estimates of global extinction 221 Box 8.3 Success Stories
Box 8.2 Challenges & Opportunities Citizen Science and Biodiversity Monitoring 239
Are We Entering the Sixth Mass Extinction? 224 Controls of Extinction Risk 240
Drivers of Extinction 226 Demographic parameters 240
Habitat loss 226 Box 8.4 Case in Point
Overexploitation 227 Why Are Frogs and Toads Croaking? 241
Invasive alien species 227 Ecological controls 242
Climate change 228 Life-history traits 243
Most threatening factors 228 Stochastic processes 245
9 Habitat Loss, Fragmentation, and Degradation 249

Habitat Loss 250
Primary drivers of habitat loss 251
Habitat Degradation
Pollution 277
277
Box 9.1 Challenges & Opportunities Desertification 283

Land Sharing versus Land Sparing 254 Erosion and sedimentation 284
Box 9.2 Success Story Metapopulations and Landscape Mosaics 285
The Rise, Fall, and Resurrection of Detroit 257 The “classic” metapopulation model 286
Habitat loss by biome 260 Source-sink dynamics 287
Conservation of metapopulations 288
The Unsustainable Use of Palm Oil 263 Box 9.4 Conservation in Practice
Future development threats 267 The Mainland-Island Metapopulation Model 289
Habitat Fragmentation 269
Biological consequences of fragmentation 270
Species-specific responses to fragmentation 276

x | Contents
10 Overexploitation 293
History of Overexploitation 294
Psychology of Overexploitation 297
Subsistence overexploitation 314
Recreational overexploitation 317
Types of Overexploitation 299 Box 10.3 Success Story
Commercial exploitation 299 The Taimen Conservation Project 322
Box 10.1 Challenges & Opportunities Theory of Sustainable Harvesting 322

No Fish Left, or Let Us Eat Fish? 303 Sustainable yield 323
Fixed quota (Q) harvesting 324
CITES: The Convention on International Trade in
Fixed effort (proportional) harvesting 325
Endangered Species of Wild Fauna and Flora 306 Limitations of MSY models 326
11 Invasive Alien Species

Overview of the Problem 330
The Population Biology of IAS 333
329
Economic impacts 349
Positive values of IAS 351
Introduction of alien species 334 Box 11.2 Challenges & Opportunities
Establishment of alien species 338 Should Species Be Judged by Their Origin,
or by What They Do? 353
Spread of invasive alien species 341
Management and Control 354
Predicting an Invasive Alien Species Wave Front 342 Box 11.3 Success Story
To Catch a Goat 355
Impacts of Invasive Alien Species 343
Impacts on biodiversity 343 Risk assessment 356
Impacts on ecosystems 348 Risk management 361
12 Climate Change 369

Anthropogenic Climate Change 370
Predicted Response to Climate Change 374
Managing Effects of Climate Change on
Biodiversity 392
Methods and models 375 Establishing refugia and safe havens 393
Current forecasts 378 Optimizing migration pathways 395
Building evolutionary resilience 396
Toward Mechanistic Climate Envelope Models 378 Box 12.3 Challenges & Opportunities
Documented Responses to Climate Change 382 Assisted Colonization: A Key Tool for Conservation,
or Pandora’s Box? 397
Population decay and local extirpation 382
Using biodiversity for climate mitigation 398
Box 12.2 Case in Point Developing green infrastructure 398
Meet the First Species to Go Extinct Because of
Anthropogenic Climate Change 383 Box 12.4 Success Story
Geographic range shifts 384 China’s Sponge City Program: Using Green Infrastructure
to Mitigate Natural Disasters 400
Altered phenologies 386
Biome regime shifts and alternative states 388
Change in ecosystem services 389

Contents | xi
Part IV Approaches to Conservation 403
13 Species-Level Conservation 405

Record-Breaking Recovery: Island Foxes of
Estimating Animal Density: The Distance Sampling
California 406 Approach 423
Goals of Species-Level Conservation 408 Capture-mark-recapture techniques 424
Challenges to Conserving Box 13.3 Case in Point
Small Populations 409 Detector Dogs Support Research and Conservation
Demographic stochasticity 410 Efforts 425
Environmental stochasticity 411 Models of Population Size 426

Loss of genetic variability 413 Geometric growth model 427
Effective Population Size 415 Exponential growth model 429
Determining effective population size 415 Logistic growth model 429
Unequal sex ratio 416 Age- and stage-based models 430
Variation in reproductive output 417 Stochastic models 433
Population fluctuations and bottlenecks 417 Predicting Population Persistence 434
Extinction Vortices 420 Minimum population size 434
Population viability analysis 436
Estimating Population Size 421
Census 421 Conservation Trade-Offs 440
14 Community and Ecosystem Conservation 445

Classification of Protected Areas 446
Strict nature reserves and wilderness areas
Effectiveness of Protected Areas
Terrestrial protected areas 461
461
(Category I) 446 Marine protected areas 463

National parks (Category II) 448
Natural monuments (Category III) 448 Innovative Partnerships Protect One of Earth’s
Habitat/species management area Last Remaining Marine Wilderness Areas 464
(Category IV) 448 Impacts on Human Well-Being 466
Protected landscape/seascape
Predictors of Success 469
(Category V) 448
Effective management 469
Managed resource protected area
(Category VI) 449 Stakeholder and community involvement 471
Integrated social development and conservation
Global Status of Protected Areas 449
goals 472
Approaches for Choosing Protected Areas 453
Hotspots of biodiversity 453
Fences and Fines … or … Integrated Conservation
Ecoregions 455 and Development? 474
Conserving Half: A Bold Agenda for Conservation 457
Political and economic boundaries 457
Ecosystem services 458
Areas of cultural importance 459

xii | Contents
15 Landscape-Scale Conservation 477

Creating Networks of Protected Areas 479
Establishing new protected areas 479
Incorporating unprotected areas into conservation
plans 497
Criteria used to select and prioritize new protected Conservation opportunities in urban habitat 499
areas 481 Conservation opportunities in agricultural
habitat 502
Conservation opportunities in mixed-use
Indigenous Protected and Conserved Areas: The Right to
Decide 482 habitat 504
Tools used to optimize selection of protected Managing the Conservation Landscape 506
areas 487 Monitoring 506
Box 15.2 Conservation in Practice Modeling 510
Decision-Making Algorithms 488 Ecosystem-based management 511
Connecting Individual Protected Areas Landscape-Scale Challenges 515
into Networks 493 BOX 15.3 Case in Point
Community-Based Natural Resource Management 516
16 Ex Situ Conservation 521

A Comprehensive Conservation Plan Saved
Resurrection Science: Should Extinct Species
the Giant Panda 523 Be Brought Back to Life? 541
Ex Situ Conservation Facilities 524 Box 16.3 Conservation in Practice
Zoos and aquariums 524 Measuring Inbreeding and Outbreeding 543
Botanical gardens 531 Reintroduction and reinforcement 544
Gene banks and seed banks 534 Box 16.4 Success Story
Contributions to In Situ Conservation 538 Reintroduction Programs That Saved Species
Captive-breeding programs 538 from Extinction 546
Public education and engagement 549
Final Thoughts on Ex Situ Conservation 551
17 Conservation and Sustainable Development 555

What Is Sustainable Development? 556
International Efforts to Achieve Sustainable
Snow Leopard Enterprises 572
Development 558 Examples of Sustainable Development Projects 573
The United Nations 558 Ecotourism and biodiversity 573
International conventions 560 Illegal wildlife trade 574
Funding for Sustainable Development 565 Box 17.3 Case in Point
The World Bank 565 Should the Ban on Elephant Ivory Trade Be Lifted? 575
Global Environment Facility 567 Fisheries 576
Individual governments 568 Climate change 577
Non-governmental organizations 570 The Future Success of Sustainable Development 578
Box 17.1 Challenges & Opportunities Box 17.4 Conservation in Practice
China’s Belt and Road Initiative 570 The Social Progress Index 582
Concluding Remarks 583
Glossary G-1
References R-1
Index I-1

Preface
Earth is home to a spectacular variety of life. Humans share the planet with more
than a million described species, and there are probably another 8 million multicellular
companions that are unknown to science, and as many as 1 trillion types of unicellular
prokaryotic organisms we have yet to discover. This amazing biodiversity is the most
striking feature of our planet, and the only thing that makes Earth unique.
But the variety of life on our planet is in trouble. Humans now dominate every eco-
system on Earth, and we have co-opted the vast majority of limited resources for our-
selves. As this book goes to press, an inferno is destroying the Amazon rainforest at an
alarming rate as illegal clearing and burning makes way for cattle ranching. So it is no
surprise that biodiversity is being lost at rates that are unprecedented in human history.
We are at a crossroads. We have just a few generations to learn how to be good stew-
ards of our natural resources, and to live sustainably alongside the other nonhuman
species that share our planet. If we do not succeed, much of the biodiversity on Earth,
perhaps the majority of it, will be lost for good.
Why did we write this book?

We wrote this book to inspire the next generation of conservation biologists to help
humans become better stewards of the world’s biodiversity. We also wrote this book to
fill a gap in the education and training of conservation biologists. While most universi-
ties have entry level classes in conservation that are taught as part of their traditional
programs of biology, ecology, and environmental science, the number of universities that
offer more advanced courses is comparatively small. It is even more rare for universities
to offer interdisciplinary courses in conservation that extend beyond the discipline’s his-
torical roots in natural sciences to include the disciplines of social sciences, engineering,
and humanities that have become central to the modern mission of conservation. The
lack of advanced, interdisciplinary training has resulted in a paucity of graduates who
have the full set of skills needed to be successful in jobs. In fact, the skillset of graduates
in some countries is so underdeveloped that many conservation organizations have had
to design their own training programs to prepare their entering workforce.
Who is this book for?

This book is for aspiring conservation biologists who seek advanced training. The book
was developed to support classes designed for upper-division undergraduates who have
already had some introduction to environmental science, ecology, wildlife biology, for-
estry, or other field related to conservation. In addition, the book was developed to
support classes designed for beginning graduate students, such as those in the growing
number of professional master’s programs that provide advanced degrees in environ-
mental science, policy, management, or sustainability.
Our text is also relevant for those who want a more interdisciplinary perspective
than is typical of introductory conservation textbooks. Conservation is no longer a
bunch of ecologists, wildlife biologists, or other natural scientists trying to save their
favorite species in a dwindling habitat. The modern practice of conservation relies
on numerous disciplines from the social sciences that account for human behaviors,
values, needs, and decision making. Modern conservation relies on disciplines from

xiv | Preface
engineering and architecture to help plan, design, and construct practical solutions
to problems. Modern conservation also relies on disciplines from the humanities that
compose law and policy, and that communicate effectively through literature, art, and
photography. Interdisciplinary approaches to conservation are integrated throughout
the text, and form the foundation for several individual chapters.
What pedagogical approaches does the book use?

To meet the conservation challenges of the future, the next generation of practitioners
will need to be better equipped than our predecessors. They will need a broader tool kit
that includes an ever-increasing body of quantitative methods and predictive models.
In addition, they will need a greater variety of qualitative tools such as stories, narra-
tives, and art that serve as effective forms of communication. This text uses a variety of
pedagogical approaches to develop the practitioner’s tool kit:
• Success Stories, included in every chapter, detail conservation biology’s most

important achievements to date. These stories provide inspiration, and remind
us that we can succeed.
• Conservation in Practice boxes detail some of the most common methods,
models, and techniques that form the quantitative foundations of the field
of conservation.
• Case in Point boxes describe case studies that reinforce key topics covered
in the chapters. They are designed to help translate concepts into real-world
applications.
• Challenges & Opportunities boxes summarize contemporary challenges,
controversies, and opportunities that future practitioners will face and need
to resolve over the course of their careers.
• Suggested Exercises provide opportunities to practice the methods, models,
and techniques described in the text, and to explore datasets and sources of
information that practitioners commonly use.
• Supplemental Videos are used to reinforce topics in each chapter with
compelling real-world examples and applications. These can be assigned with
readings, or used to augment lectures and discussions.
• Chapter Summaries succinctly recap the take-home points of each chapter.
• For Discussion questions can be assigned, or used to facilitate in-class
discussions that get students thinking more deeply about the key points and
issues from each chapter.
• Suggested Readings provide a list of additional papers students can read,
or that instructors can assign, if there is a desire to dig deeper.
In addition to the pedagogical approaches, students are provided with a companion

website that organizes all supplemental online material and keeps the weblinks current.
Instructors resources are also available online to aid instructors in creating presentations
and to assess student understanding.
What is our hope for this book?

It took us a long two years to develop the first edition of this textbook, and there were
many ups and downs along the way. But we were determined to complete the project
because of our optimism that the current trajectory of biodiversity loss can be altered
for the better. We need a new generation of individuals who have both a passion for
nature’s living things, and the skills needed to help conservation succeed more often
than it has in the past. For those who have both the passion and the skills, we believe
you will have a rewarding career filled with purpose. Our sincere hope is that this book
will help you become the next great champion for biodiversity.

Acknowledgments | xv
Acknowledgments
We want to thank the large number of individuals who provided us with data, figures,
and tables that are used throughout this textbook. The next generation of practitioners
will stand atop your work. We extend our sincere gratitude to the many individuals who
graciously reviewed various chapters to help us improve this book. This includes several
anonymous reviewers, as well as the following outstanding individuals:
Dr. Matthew S. Becker, Prof. Aaron M. Ellison, Heidi Marcum, PhD,

Zambian Carnivore Programme Harvard University Baylor University
Andrew F. Bennett, Joel T. Heinen, Stephen G. Mech,
La Trobe University Florida International University Albright College
Nate Bickford, Ray Hilborn, Curt Meine,
University of Nebraska, Kearney University of Washington University of Wisconsin, Madison
Robert Boyd, Joshua D. Holbrook, Ben A. Minteer,
Auburn University Montreat College Arizona State University
Joseph W. Bull, Vanessa Hull, Vincent Nijman,
University of Kent University of Florida Oxford Brookes University
Abigail E. Cahill, Daniel Karp, Liba Pejchar,
Albion College University of California, Davis Colorado State University
J. Baird Callicott, Christopher Kellner, PhD, Charles Perrings,
University of North Texas Arkansas Tech University Arizona State University
Scott Connelly, Richard K. Kessler, Andrew Rassweiler,
University of Georgia Campbellsville University Florida State University
Brian L. Cypher, Jason A. Koontz, Michael Reed,
California State University, Stanislaus Augustana College Tufts University
Laura E. DeWald, Jay T. Lennon, Dov F. Sax,
Western Carolina University Indiana University Brown University
Dr. Nicole Duplaix, Mark Manteuffel, PhD, D. Alexander Wait,
Oregon State University Washington University Missouri State University
We are indebted to the team at Sinauer Associates/Oxford University Press that

included Dean Scudder (President, Sinauer Associates), Martha Lorantos (Senior
Production Editor), Peter Lacey (Senior Production Editor, Media), Meg Britton Clark
(Production Specialist), Joan Gemme (Production Manager), Mark Siddall (Photo Re-
searcher), Michele Beckta (Permissions Supervisor), Tracy Marton (Senior Production
Editor), artist Jan Troutt, and copy editors Lou Doucette and Carol Wigg. You took
our imperfect text and partially developed ideas and turned them into a polished and
beautifully illustrated book that we all can feel proud of. Thank you!
Lastly, we thank our families for putting up with us for two years as we completed
this project. Your love and patience were inspirational, and we were touched.
Bradley J. Cardinale
Richard B. Primack
James D. Murdoch
August 2019

Media & Supplements
to accompany
oup.com/he/cardinale1e
For the Student For the Instructor
The Companion Website for Conservation Biology, (Instructor resources are available to adopting
First Edition, provides students with study and instructors online. Registration is required. Please
review tools to help them master the material contact your Oxford University Press representative
presented in the textbook, all free of charge and to request access.)
requiring no access code.
The Instructor Resources for Conservation Biology,
The site includes the following resources: First Edition, include a variety of tools to help
instructors incorporate visual resources into their
Videos and Audio – Engaging media clips
lectures and course materials and enhance student
provide information and inspiration on a range
learning. The site includes:
of conservation biology topics.
Suggested Exercises– Students can learn Figure JPGs– Includes all of the figures and
essential skills and practice analyzing real data tables from the textbook, formatted for optimal
with these thoughtful exercises. legibility when projected. Complex images are
provided in both whole and split versions.
Web Links– Connect with the many
organizations and institutions discussed in the Figure PowerPoint Slides– Features all the
text with helpful web links. figures and tables from each chapter, with titles
on each slide, and complete captions in the
Chapter Outlines– Useful study aids summarize
“notes” field.
each chapter’s key concepts.
Suggested Exercise Resources– For many of
Flashcards– Students can gain mastery over each
the Suggested Exercises, instructors have access
chapter’s key vocabulary and concepts with this
to helpful materials, such as preformatted
helpful flashcard set.
spreadsheets for data-analysis exercises and
answers to exercise assessments.
eBook (ISBN 978-1-60535-882-6)

Conservation Biology, First Edition, is available as an eBook via several different eBook
providers, including RedShelf and VitalSource. Please visit the Oxford University Press
website at oup.com/ushe for more information.


Part I
Foundations of
01_Cardinale1e_CH1.indd 1 8/26/19 12:50 PM

1
The State of
Our Planet
© Katesalin Pagkaihang/Shutterstock.com
B
y‌ many if not most measures, humans are the most
remarkably successful species to ever inhabit planet
Earth. We have evolved one of the largest brains per
body size that exists in the animal kingdom, which has given
us an unparalleled intellectual capacity and ability to reason.
Our species is one of the more social organisms on the planet
(particularly among mammals), which has led to cooperation,
advanced learning, and a division of labor that has fostered
our success. Our mastery of tools and our technological
advances are unparalleled among other life-forms. These, and
other qualities, have allowed our species population to grow
rapidly and to conquer and dominate more of the planet than
almost any previous organism.
But there is one measure that humans do not yet excel
in, compared with other species—longevity. Homo sapiens
have existed on this planet for a mere 200,000 years, which is
but a fraction of the time that most species survive on Earth.
Will humans be able to sustain themselves for another million
years and achieve the longevity that is typical for one of Earth’s
species? Many of the very qualities that have allowed us to
be so remarkably successful thus far are eroding the natural
infrastructure of the planet—the collection of genes, spe-
Orangutans are presently found only in the rain
cies, and biological communities that compose the natural and forests of Borneo and Sumatra, where they are
managed ecosystems on which people depend. threatened with extinction by deforestation.

4 | Chapter 1
If we are to ultimately become one of longest-lived species in Earth’s history,

humans are going to have to use our intellect to learn how to live more sustain-
ably on finite resources. We are going to have to use our tools and technology to
conserve the biophysical processes that regulate life on the planet. And we will
have to do so while using our social cooperation to generate a decent quality
of life for our fellow humans. If we succeed, then Homo sapiens will indeed be-
come the most remarkable species by all measures. If we fail, then our era will
be recorded in the geological record as one of the greatest periods of biological
loss in the history of life on Earth.
This chapter gives an overview of the current state of biology on our planet.
It is the precursor to all other chapters in this book, as it establishes some ba-
sic facts about the forms of environmental change that are occurring across
the globe as a result of human domination of the planet and the subsequent
impact that these changes are having on biodiversity. We begin with the state
of the human species, emphasizing gains that have allowed our species to be
successful, while also emphasizing the social inequalities that are preventing
many from having a good quality of life. We then review the state of our global
environment, emphasizing how human gains for some have come at a cost to
the environment. This, in turn, leads us to overview the state of biodiversity,
including biodiversity losses that are occurring as a result of environmental
change. We end the chapter with some reasons to be optimistic that we can, in
fact, become the most remarkable species ever to inhabit planet Earth—a spe-
cies that can provide a high quality of life to our own kind while, at the same
time, protecting the myriad of amazing life-forms that share our planet.
State of the Human Species

Modern humans first appeared in the fossil record about 200,000 years ago,
which is but a tiny fraction of the 3.5-billion-year history of life on Earth (Figure
1.1). For most of our brief history, our ancestors lived as hunters and gatherers—
a dangerous way of life that kept population sizes small, probably less than 10
million individuals across the entire globe.
As humans transformed from hunter-gatherer to agrarian societies, com-
munities evolved that could support more people. By roughly 1 ce, the world
population had expanded to 300 million, and it continued to increase at a con-
stant and steady rate for hundreds of years. Then the Industrial Revolution hit.
Living standards in the eighteenth century rose dramatically in the developing
countries of Europe and in the United States, and improvements in technology
and medicine reduced the prevalence of famines and epidemics in many parts
of the world. Population growth accelerated quickly, and humans reached their
first 1 billion individuals about 1800.
While it took nearly 200,000 years for Homo sapiens to reach the first 1 bil-
lion, it took just 130 years to reach the second billion, 30 years to reach the third
billion, and 15 years to reach the fourth billion. In a span of less than 200 years,
our species increased more than sevenfold to the current mark of 7.6 billion
Video 1.1 See the staggering in 2017 (see Figure 1.1). This staggering rate of growth is almost impossible
▲
growth of the human population to convey with graphs or words (Video 1.1). No large vertebrate animal in the
over time. history of the planet has ever grown as quickly, and with as much success, as
oup-arc.com/e/cardinale-v1.1 Homo sapiens, and that is a testament to our unique intellect, innovation, and
technological advances as a species.
The human population recently passed an inflection point called the demo-
graphic transition, where global growth has now shifted from unconstrained,

The State of Our Planet | 5
Figure 1.1 Human dominance is but a small fraction of Earth's 10
World population growth (in billions)

4.6-billion-year history. Although Homo sapiens evolved 200,000 years
ago, nearly all of human population growth has occurred in the last 8 Growth is slowing
200 years. Now at 7.6 billion, the population has passed an inflection
point (red dot) where growth has slowed and is heading toward a 6
steady state. (After T. P. Soubbotina et al. 2000. Beyond Economic
Growth: Meeting the Challenges of Global Development. Washington, 4
DC: World Bank.)
Developing Industrialized
2 regions
regions
0
1750 1800 1850 1900 1950 2000 2050
Year
Years (in billions)
Hadean Archean Proterozoic Phanerozoic
4.56 4.0 3.5 3.0 2.5 2.0 1.5 1.0 0.5 0
mrowka; Mike Peel/CC BY-SA 2.0; © CHOTE

© iStock.com/Naeblys; © iStock.com/dzika_
Wisconsin-Madison; spot photos from left to

right: © Diego Barucco/Shutterstock.com;
Present
Timeline by Andrée Valley, University of

Formation Earth’s Moon Earliest Earliest Rise in First First Dinosaurs Humans
of Earth core formation isotopic fossils atmospheric cells with hard-shelled
BKK/Shutterstock.com
forms evidence (Apex oxygen nucleus animals
for life Chert)
Evidence
for oceans
exponential increases to reduced logistic growth that is headed toward a steady

state (see Figure 1.1). This inflection point represents the transition from high
birth and death rates to lower birth and death rates that have occurred as countries
have transitioned from preindustrial to industrialized economic systems. There
are many potential triggers of the demographic transition. Social triggers include
reduced fertility rates caused by improved access by the population to contracep-
tion, a reduction in subsistence agri-
culture, an increase in the status and
education of women, and a reduction
© Rawpixel.com/Shutterstock.com
1000
in the value of children’s work.1 Eco-
logical triggers include limitation by
Maximum population (billions)
essential resources like energy, food,

or water that become increasingly
scarce as the population approaches 100
Earth’s carrying capacity. 2,3
Whatever the triggers—whether Upper interval

social, ecological, or both—it is now = 12 billion
Conservation Biology 1e Cardinale
clearOxford
that University
growth of the global hu-
Press 10
manTroutt
population has slowed and Lower interval
Visual Services
= 7.7 billion
will Cardin_01.01_v2.ai
soon reach some type of steady
8-20-19
state, perhaps as early as the middle
of this century. There have been at 1
least 65 attempts to estimate Earth’s 1650 1700 1750 1800 1850 1900 1950 2000
Year
carrying capacity for humans (Fig-
ure 1.2). While these range from ex- Figure 1.2 This graph shows 65 estimates of Earth’s carrying capacity for humans
treme pessimism (1 billion) to un- (y-axis) that have been published through time (x-axis). The credible interval for these
constrained optimism (1000 billion), estimates ranges from 7.7 to 12 billion, closely matching projections from the United
the credible interval of the estimates Nations of 7.8 to 12.5 billion. (After J. E. Cohen. 1995. Science 269: 341–346.)

6 | Chapter 1
ranges from 7.7 to 12 billion,4 which closely matches projections from the United
Nations. Given that the human population is already at the lower bound of these
estimates, and given that the world’s current growth rate of 1.14% has a potential
doubling time of 61 years, those reading this book will likely be the first people in
history to know what Earth’s carrying capacity actually is for humans.
Given the staggering growth of the human population, it is perhaps surprising
that the “average” person born in the modern era will live a longer, healthier, and
more prosperous life than the average person born in any other period of human
history. Mortality rates in premodern countries for children under the age of five
were routinely 300 to 500 deaths per 1000 live births.5 But during the twentieth
century, modern medicine, central heating, improved hygiene, cleaner water, and
access to more food have all helped reduce infant mortality to historically low
rates (Figure 1.3A).6 Those born in the premodern world had life expectancies of
a mere 30 years.5 Since 1900 the global average life expectancy has more than
doubled, exceeding 76 years at the start of the twenty-first century. The average
person in the modern era will also have a better standard of living compared with
his or her ancestors. Since the 1800s, the global gross domestic product (GDP)
per capita has increased more than tenfold (Figure 1.3B).7 Modern technology and
the green revolution have led to nearly a doubling of per capita caloric intake
in modernized countries,8 and the per capita calorie supply has been increasing
steadily in nearly all regions of the globe since the 1960s.9
While conditions have improved for the “average” person on the planet, the
benefits of economic development have been unequally distributed across the
globe, leading to vast inequalities among countries and among those living in
individual countries (Video 1.2). For example, although Americans constitute
Video 1.2 Consider the just 5% of the world’s population, they consume 24% of the world’s energy.10,11
▲
argument for a more equitable If everyone on the planet consumed as much energy, food, water, minerals,
future for humanity. and space as the average U.S. citizen, we would need four Earths to sustain
oup-arc.com/e/cardinale-v1.2 them. There are also huge disparities among individuals. The wealthiest 10%
of the world’s population accounts for 59% of all consumption on the planet,
(B)
(A) 8000
2008
175 80 7000
Infant mortality ( per 1,000 births)
165 Life expectancy

(1990 international dollars)
Life expectancy at birth ( years)
150 76 6000
World GDP per capita
60
125 5000
47 Life expectancy
100 4000 GDP has been
increases while infant
© iStock.com/Sezeryadigar
40 rising exponentially.
75 morality rates drop. 3000
Infant
50 2000
mortality 20
25 1000
7
0 0 0
1900 1950 2000 1500 1600 1700 1800 1900 2000
Year Year
Figure 1.3 The average child born in the modern era will have 1900–1970, U.S. Public Health Service, Vital Statistics of the
(A) a greater chance of survival and a higher life expectancy United States, Vols. 1 and 2; 1971–1997, U.S. National Center for
and (B) greater per capita wealth than any generation in human Health Statistics, Vital Statistics of the United States; National Vital
history. The resulting population growth and consumption, Statistics Report [NVSR] [formerly Monthly Vital Statistics Report];
particularly in fully developed countries, will be a major challenge and unpublished data; B, data from A. Maddison. 2009. Statistics
for conservation efforts. (A, data from U.S. Census Bureau, on World Population, GDP and Per Capita GDP, 1–2008 AD.
Statistical Abstract of the United States. 1999. Compiled from © Angus Maddison/University of Groningen.)

The State of Our Planet | 7
© dbimages/Alamy Stock Photo

Figure 1.4 While the “average” person born in the modern era will live a longer, healthier, and more prosperous
life than his or her ancestors, more than 2 billion people still live in extreme poverty and are malnourished. People
whose basic needs are not met do not have the luxury of caring about biodiversity loss. Therefore, bringing com-
munities out of poverty, particularly in developing nations, will be a major challenge for conservation efforts.
whereas 1.3 billion people continue to live in absolute poverty, surviving on less
than US$1.25 per day (Figure 1.4).12 Still another billion people live in sufficient
poverty that they are malnourished, with food shortages disproportionately
represented in less-developed countries of the world. While the number of poor
and malnourished people is lower than recorded a few decades ago (suggest-
ing that some progress has been made), the need to alleviate the suffering of
those living in extreme poverty is still considered to be the greatest challenge
to achieving a sustainable human population on the planet.
Suggested Exercise 1.1 Use the Global Footprint Network’s website to

estimate your ecological footprint, which is the number of planets we’d need
if everyone lived like you. oup-arc.com/e/cardinale-ex1.1
State of the Global Environment

With our vastly expanded population size, increased affluence, and improved
technologies, human activities now control almost all physical and biological
processes that regulate life on the planet. While we do not wish to present a
“doom-and-gloom” narrative in this book, it is necessary to establish some basic
facts at the beginning of this book about what is happening across the globe as
a result of human domination of the planet. These facts form the template for
actions to save biodiversity.
Land
More than 50% of the inhabitable land surface of the planet has now been
transformed in some way to support the growing human population.13 To date,
5 billion hectares (50 million km 2 ) of natural habitat have been converted to
croplands and pastures,14 and an additional 350 million hectares (3.5 million
km2 ) of the world’s land surface have been converted into urban environments.15

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Crœsus’ succession to the great dominion which Alyattes had left
was not undisputed. But the son had inherited the vigour of his
father. He anticipated the plans of his rival. The pretender
disappeared,—how or whither is not known; and his supporters, who
were largely drawn from the feudal nobility of the land, met death in
many grievous forms. Some of the Greek cities had more than
sympathized with his antagonist, so to these he now turned his
attention. All of them, Æolian, Ionian, and Dorian alike, were reduced
to the position of Lydian dependencies, though in matters purely
local they remained autonomous, if the name of autonomy can be
given to a form of government in which a local tyrant played the part
of administrator and political agent. Yet unpromising as was their
position from the point of view of theoretical politics, they were in
actual fact treated with marked consideration by Crœsus.
It is impossible at the present day to sound the motives which
underlay the attitude which this extraordinary man adopted to the
Greeks alike of Asia and of Europe. It may have been from pure self-
interest; it may have been because Hellenism had cast over him the
glamour which it cast over other barbarian monarchs. Whatever the
cause, the fact remains that, when once he had reduced the Greek
cities to that position of dependence which was necessary for the
political homogeneity of his empire, he seems to have lost no
opportunity of ingratiating himself in the eyes of his Greek subjects
and of their kinsmen beyond the seas. At Branchidæ and at Ephesus
he enriched the Greek temples with splendid offerings; and the
wealth of the gifts he gave to Delphi excited the admiration of
centuries. If contemporary report be not exaggerated, the value of
his dedications to the great Hellenic shrine amounted to considerably
more than a million pounds sterling of the money of the present day.
Gifts of great value were also sent to the lesser Greek oracles.
The authorities at Delphi would have been
DESIGNS OF
exceptional among similar societies in all ages
CRŒUS.
of the world, had they not shown appreciation
of a devotee so wealthy and so willing. He was made a citizen; to his
embassies were given a precedence over all others.
It is difficult to imagine that Crœsus should have expended such
enormous sums on the cultivation of friendly relations with the
Greeks of Europe for purely sentimental reasons. The oracles were
not the only recipients of his gifts. The friendship of prominent and
powerful families in various States, such as the Alkmæonidæ of
Athens, was bought with a price.
Perhaps the explanation maybe sought and found in the previous
and later policy of the king. He had subjugated the Greek cities of
the coast, and by so doing had advanced his kingdom to its extreme
limits on the west. Unless he converted Lydia into a naval power,
further expansion on this side was impossible. So he turned his eyes
towards the East, where nature and the circumstances of the time
offered what must have seemed a favourable opportunity for the
extension of his empire.
It must, however, have been quite evident to him that any policy of
expansion eastward could only be carried out with safety in case his
rear was secured from attack, where danger lay, not merely in the
recently subdued Greek cities, but also in the possibility of any
movement on their part being supported by help from their kinsmen
in Europe.
Considerations such as these must have had a large influence
upon his policy.
The story of his operations in the East has survived in history in
what is manifestly a very mutilated form. It is fortunate that Strabo
has preserved some reliable details which Herodotus does not
mention in his somewhat legendary account of the last days of the
rule of Crœsus. The Lydian frontier had been extended to the Halys;
but the motley collection of races and states included within the
dominion was in some cases bound to the ruler of Sardes by
comparatively loose ties. These ties Crœsus strengthened.
Affairs in Asia beyond the Halys were at the moment, when
Crœsus brought his plans to maturity, about the year 548, in a
condition which made all certain calculation as to their issue
impossible. The Median dynasty had come to an end some four
years before, and with it the treaty concluded by Lydia with the Mede
in 585 had come to an end also. Cyrus must have been an unknown
factor to the Lydian, though doubtless the merchant travellers had
brought back from the East many a tale of his energy and success.
He was certainly a danger: and the question probably suggested
itself to Crœsus whether he were not a danger which it would be
wise to forestall, by pushing forward the Lydian frontier to that mass
of mountains formed by the meeting of Taurus with the Armenian
chains. Such a precautionary measure would be rendered the more
attractive to the Lydian trader by the fact that it would lead to the
inclusion within the empire of that rich mineral district on the south
shore of the Euxine wherein the famed Chalybes dwelt.
Crœsus was wise enough not to enter upon this venture single-
handed.
It is evident that the comparative indifference with which
Nabonidus and the Babylonians had originally regarded the change
of rulers in Median empire, had by this time given place to a feeling
of uneasiness, if not of actual alarm. The easy-going, peace-loving
antiquarian of Babylon might well be apprehensive as to what might
be the next object of the uncomfortable enterprise of his energetic
neighbour. Even then the faint outlines of the writing on the wall were
well-nigh decipherable.
Amasis of Egypt had far less grounds for alarm; but even he
seems to have caught the infection of fear.
H. i. 77.
With these two states Crœsus entered into
negotiations, which resulted in the formation of a
grand alliance, having for its object the suppression of the power
which was so rapidly developing in the East.
The negotiations of Crœsus were not confined to the great
powers. He sought and obtained allies in European Greece. The
Lydian kings had had a long experience of the value of the Greek
heavy-armed infantryman. Greek hoplites had fought many a time
both with and against them. The addition of a
LYDIA AND SPARTA.
contingent of them to the grand army which the
king was now gathering together would be of inestimable value.
There was evidently a difficulty about his obtaining such a force from
the Greek cities of Asia; nor can there be any reasonable doubt as to
where that difficulty lay. These cities had within the last few years
been robbed of much of that measure of autonomy which they had
up to that time enjoyed, and upon which they had set a value out of
proportion, doubtless, to its real worth. The vivid discontent which
such a loss must have aroused in Greek minds, a discontent the
depth of which the experience of ages would enable the Lydian to
gauge, would inevitably render them dangerous elements in a Lydian
army. The cities did, indeed, with one exception, remain proof
against Cyrus’ attempts to tamper with their loyalty; but their attitude
at the time was probably as much due to caution as to fidelity. Their
geographical position would not allow them to accept risks against
Lydia.
It was, therefore, to Greece itself that Crœsus turned. The
relations which he had so assiduously cultivated with Delphi enabled
H. i. 69.
him to obtain its assistance in the negotiations. The
outcome was, so Herodotus says, that Sparta, partly
persuaded by the oracle, partly flattered by the Lydian embassy,
consented to give aid in the war. Moreover, the way to this alliance
had been previously paved with Lydian gold.
It is true that this contingent never reached Lydia. Ere it had
actually started, Sardes had fallen and Crœsus was either dead or a
prisoner. Whether the delay in despatching it was intentional or not,
the satisfactorily attested fact of such an alliance having been made
is evidence that the Lydia of that day exercised a very real influence
in Greece. Of the danger to which Hellenic civilization was exposed
by Lydian friendship, enough has been already said.
That friendship was genuine and unaffected on the side, at any
rate, of the Greeks. The relations of Crœsus with Delphi must have
been largely instrumental in forming it; but what happened in relation
to this very war showed clearly that the feeling of Greece towards
Crœsus was built upon wider foundations. The Greeks had come to
regard him as a distinguished convert to that Hellenism they so
much loved. The impression may have been false, but it was
powerful. “He loveth our nation” is an article in a national creed
whose possibilities can be hardly exaggerated. That the feeling had
become independent of the relations with Delphi is conspicuously
shown in the present instance by the fact that it was Delphi which
administered to it a shock which the Greek world took long to forget.
The remembrance of it was evidently vivid a hundred years later in
the time of Herodotus.
It came about as follows. Anxious as to the issue of the great
venture upon which he was entering, Crœsus sought to fortify or
defeat his own resolution by inquiring of the oracles as to what the
future had in store for him.
H. i. 53.
Two of the oracles consulted, of which Delphi was
one, answered that “if he warred with the Persians he
would overthrow a mighty empire.” The response was capable of two
interpretations, of which Crœsus seized upon the most obvious; and
was thus, so the Greeks thought, led to his undoing. Despite the
pious faith with which Herodotus regards the utterances of the
Delphic oracle, he is unable to conceal the tremendous shock which
this apparent deception caused to Hellenic sentiment all the world
over. To the Greek it appeared as though the oracle had betrayed its
best friend and his also. Even in the cities of Asia, chafing though
they were under recent subjugation, this feeling must have found
some echo, whose resonance lasted till Herodotus’ own time. It is
unlikely that he would ever have disclosed its existence had not the
feeling been very widespread in the Hellenic world he loved. The
H. i. 90.
legendary story which he relates of the conversation
between Crœsus and Cyrus, expresses evidently a
feeling entertained by many besides Crœsus himself; and in the
chapter which follows upon this tale, he shows that the Delphic
oracle was forced by public opinion to attempt to explain away the
apparent deception it had practised. The true
FALL OF THE
LYDIAN KINGDOM.
explanation, which would have relieved it of a
large part of the burden of the moral guilt, was
one it dare not give in view of the prophetic character which it had to
maintain before the eyes of the world. Prophecy founded upon
intimate knowledge of Greek affairs was very far from being the
mere guesswork, wrapped in enigma, of its utterances relative to
matters deep in Asia, of which it can have had no real ken.
The account of the campaign given by Herodotus is full of
inconsistencies; but by comparison of his story with other incidental
references to it in various sources, it is possible to arrive at an
understanding of the main outlines of what took place.
The great coalition might have taken Cyrus by surprise, had not
the plans of Crœsus been divulged to him by an Ephesian traitor, if a
tale preserved by Diodorus is to be believed. The mere fact that he
was able to anticipate the designs of Crœsus renders it probable that
some disclosure of the kind did take place.
Forewarned and forearmed, Cyrus executed a rapid and
adventurous march through the northern territories of the Babylonian
kingdom, and must have been already near the Taurus before
Crœsus received from his ally Nabonidus news of the coming attack.
He was but half-prepared; but the danger was so imminent that he
had to take the field with the force he had with him, while he sent
urgent messages to his allies to come with all speed to help him. He
H. i. 75.
crossed the Halys into the district of Pteria, which he
laid waste as a defensive measure. The historians,
Herodotus and Polyænus, are hopelessly at variance as to what
happened in the actual fighting that ensued. A great battle did take
place: that is certain. It is also certain that after the battle Crœsus
retired through Phrygia to Sardes; but whether he did so because he
had been defeated, or because he had inflicted a severe check on
Cyrus, and expected that a diversion on the part of the Babylonians
would make it impossible for him to advance towards Sardes in the
winter, is unknown. In any case, Nabonidus did not move, and Cyrus
surprised Crœsus in Lydia. Crœsus, caught unprepared, made a
desperate defence with such forces as he could collect; but he was
shut up in Sardes. Of the real history of the siege the Greeks seem
to have known little or nothing; their chroniclers give the most
contradictory accounts of it. But the town fell within a short time—
taken, it would seem, by escalade. What became of Crœsus is not
Bakchylides
known. It is probable that he immolated himself upon
III. 23 ff. a burning pyre. The tale was too shocking for Greek
ears, and was softened down by a legendary addition
to the effect that he was saved from the flames by divine
intervention.
The sudden collapse of Lydia is one of the most remarkable
incidents in history.
It fell in a moment, as it were, never to rise again; and it fell, not in
the decadence of age, but at the very height of its young and
vigorous life. To the Greek the spectacle was bewildering: nor is it
strange that a catastrophe so sudden and complete, unparalleled,
indeed, in the history of the world, should have so dazed the senses
of those who were spectators of it, that they were never able to give
a rational account of how it came to pass.
CHAPTER II.
PERSIAN AND GREEK IN ASIA. THE SCYTHIAN
EXPEDITION.
Despite the great catastrophe which had just taken place before
their eyes, the Greek cities had no mind to make an unconditional
surrender to the power which had vanquished their old master. It was
unfortunate that, after coming to such a decision, they did not
combine in a common resistance. The inherent weakness of their
strategical position, together with the incompleteness of the
sympathy between Æolian, Ionian, and Dorian Greek, made such
united action difficult. There is a terrible sameness in the drama of
history as played upon this coast of the Ægean. The scenery
admitted of but one plot, of which the leading motive was disunion. In
the present, as in other instances, the Dorian states of the Carian
coast went their own way. They threw in their lot with their Carian
neighbours. The Æolians and Ionians were not altogether blameless
in the matter. They did not at first show a bold front to the Persian,
but offered to submit to him on the terms on which they had
submitted to Crœsus.
Save in the case of Miletus, the largest and most formidable of
the towns, Cyrus would not hear of terms; and so the cities prepared
to fight for their liberty.
It was the beginning of the winter; and, as the Persians did not
possess the means for assaulting the cities from the side of the sea,
the latter had a few months’ respite wherein to make preparations.
They appealed to Sparta for help. That cautious government, which
was probably congratulating itself on having escaped from involving
its citizens in the Lydian débâcle, refused active assistance, but sent
an embassy to Cyrus to warn him against interference. Cyrus, whose
notions as to the geography of this part of the world may well have
been vague, asked who the ambassadors were, and whence they
H. i. 153.
came. On being told, he warned them that, all well, he
would give them cause to talk about their own woes
and not those of the Ionians. This rough humour must have seemed
in great contrast to the politeness with which Crœsus had addressed
the foremost race in Greece.
Cyrus was obliged to entrust the completion of the conquest of
the Lydian kingdom to one of his lieutenants. The news of the fall of
Sardes had scared Babylon into inactivity; but the Baktrians and
Sakæ on the extreme eastern borders of his dominion had seized
the opportunity afforded by the western campaign to rise in revolt.
He had not proceeded far on his homeward march before news
reached him of a rising in Lydia. Paktyas, a renegade Lydian who
had embraced his cause, and to whom the conqueror had entrusted
the care of the transport of the spoils, had intrigued with the Ionian
Greeks; and, having ample funds at his disposal, had hired
mercenaries from them. Tabalos, the Persian lieutenant whom Cyrus
had left behind him, was besieged in the citadel of Sardes; and there
was every prospect that, if the place fell, all the work of the late
campaign would have to be done again. There was no time to be
lost: nor was Cyrus the man to lose time. He despatched an army
under Mazares the Mede to rescue the besieged, and Sardes was
saved. Paktyas fled to Kyme, and thence to the islands. He neither
deserved nor received sympathy, and, after various adventures, was
handed over by the Chians to the Median commander.
With the flight of Paktyas the insurrection in Lydia came to an end;
in fact, in so far as extant records go, the Lydians themselves played
but little part in it. The passive and entire submission of this people,
their acceptance, once and for all, of the yoke laid upon them, is one
of the most extraordinary features of this extraordinary time. It might
well have been expected that a nation with a
CONQUEST OF
IONIA BY PERSIA.
past so recent and so glorious would have
seized the first and every opportunity of
attempting to regain the freedom, if not the dominion they had lost.
But nothing of the kind took place; and even the great effort of the
Ionian revolt failed to rouse them from the apathy of defeat.
The circumstances of Paktyas’ rebellion showed Mazares that the
Greek cities of the coast could no longer be left in a position to be
the instruments of trouble in the newly-won territory. To them,
accordingly, he immediately turned his attention. He first attacked
Priene and sacked it; but, before he had completed the reduction of
Magnesia on the Mæander, he died, and Harpagos, who succeeded
him as governor, took up the task of reduction. Phokæa and Teos
were besieged. Ere they fell, the mass of their inhabitants went into
voluntary exile—the Phokæans to Corsica in the farthest west, the
Teans to the near coast of Thrace.
There can be little doubt that the departure of these peoples was
a disaster of the first magnitude to the Greek towns of Asia. It is
hardly possible to realize at the present day the strength of
resolution which prompted the Phokæans to undertake their long
and perilous journey. They are the New Englanders of the sixth
century before Christ. Their presence fifty years later, at the time of
the revolt, might have given the Ionian resistance that “stiffening”
which it seems to have lacked; indeed, a member of the remnant
they left behind them, that dare-devil old pirate Dionysius, is the one
prominent person on the Greek side in that distressful time whom
later historians consented to praise. One by one the other Ionian and
Æolian cities fell into Persian power. There does not appear to have
been any real combined resistance. Nature had made them units
without unity. The islanders of Samos alone escaped subjection.
Caria was next attacked. It yielded practically without a blow, and
the Dorian colonies fell with it into Persian hands—a fate in their
case not wholly undeserved. Lycia fought for its liberty, but in vain;
and with its subjection the establishment of Persian rule on the
continent of West Asia was complete.
The rest of the career of Cyrus, important though it is, has little
influence on Greek history. His campaign in the East was a
prolonged one. He seems to have extended the borders of his
empire to the Thian-Shan and Suleiman ranges, if not into the plains
of India itself. His aim can hardly have been the mere acquisition of
these enormous areas of comparatively unproductive territory. The
reason lying behind his policy was, in all probability, the fact that the
races of this region were near akin to his own, and that he wished to
advance against the Semitic peoples at the head of a forced coalition
of the Iranian races.
The turn of Babylon for attack was soon to come. Nabonidus’
antiquarian researches absorbed more and more of his time, and the
real conduct of the government seems to have passed into the
hands of his son Bel-sharuzar, the Belshazzar of the Book of Daniel.
In the final struggle, indeed, Nabonidus seems to have come to the
front again.
By 538 Cyrus was ready. He made great preparations for the
invasion, probably in anticipation of a much harder task than it
actually proved. The collapse seems to have been rapid, so that
within a short time Babylon was taken, Nabonidus a prisoner, and
the brief revival of the Chaldæan empire at an end. The whole of the
Babylonian dominions submitted to the conqueror, and the empire of
Cyrus now stretched unbroken from the Ægean and Mediterranean
on the west to the borders of India on the east.
Of the three rival kingdoms, Egypt alone survived. Doubtless
Cyrus had designs upon it; but after the fall of Babylon in 538, he
seems to have wisely devoted himself to the task of consolidating
the empire he had won so rapidly in the previous fourteen years.
Before his plans were ripe for an expedition beyond the Isthmus of
Suez, disturbances in the far east called him thither. There he died,
probably in a great battle about the year 529. The halo of legend
which rapidly formed about his great personality concealed not
merely the real man, but even his real history.
THE PERSIAN
Four versions of the story of his death, each
CIVILIZATION.
differing wholly from the other, were known to
the Greeks. But whatever fate he met, his body was brought to his
home-land, where the remains of his tomb may be seen at the
present day. “I am Cyrus, the king, the Achæmenian,” is the only part
of his epitaph which survives. It would be too little for a lesser man. It
is sufficient for him even now that he has been twenty-five centuries
dead.
The new Orientalism with which the Asian Greek was brought into
contact by the conquest of Lydia was, in many respects, of a
different character to that which had preceded it in Western Asia. As
years rolled on, and the specially Persian characteristics of it
became more and more merged in the general Oriental type, the
difference tended to disappear; but even until quite late times the
hardy races from the mountains of Iran had many national customs
which were in strong contrast to the typical civilization of the
Euphrates plain. Though far from ideal, there were certain grand
elements in it which struck the imagination of some of the finer minds
of Greece, and which, through them, must have influenced Greek
life, though in ways which it is not possible now to trace. Had the
Greek come much under its influence, that influence, though it would
have been disastrous in many respects, would not have tended
wholly for evil.
The civilization was indeed essentially of an Eastern type. It is
unnecessary to point out the significance of such a general
characteristic. The Mede and Persian had been for centuries next-
door neighbours of the population of the great plains, and it was
inevitable that they should have borrowed from their brilliant life. Yet,
despite their nearness, there was a triple gulf between the two,
which the intercourse even of centuries could not bridge. Difference
of race, difference of habitat, and, above all, difference of religion
sundered them. The Iranian and Semite regarded the world and life
in it from different points of view. The struggle for existence
presented itself in wholly different aspects to the mountaineer and
the man of the plain. The monotheist could have but little sympathy
with a polytheistic creed.
The Medo-Persian was a strange product for an Asiatic soil. He
was an Asian apart. His religious belief was alone calculated to
make him remarkable among his contemporaries. The Asiatic of this
time had a natural tendency towards polytheism. The monotheism of
even the Israelites was spasmodic. But with the Persian monotheism
was the set religion of the race. It had a legendary origin in the
teachings of Zarathustra, or Zoroaster, as he appears in Western
history. Ahura-mazda was the one god. There were, indeed, other
objects of worship,—the stars, the sun, the moon, and fire, beautiful
and incomprehensible works of Ahura-mazda; but he was god alone.
Other spiritual beings there were too, represented as deified virtues
and blessings—Good Thought, Perfect Holiness, Good Government,
Meek Piety, Health, and Immortality; and these stood nearest to
Ahura-mazda’s throne.
The national religion had not, indeed, wholly escaped the
contamination of the less spiritual cults of the neighbouring peoples.
The animalism of the worship of the Babylonian goddess Mylitta had
been introduced into the land under the guise of the adoration of the
nymph Anahita.
Nevertheless, with the Persian the deification of the various forms
of nature took a special form. The deities themselves were treated
as demi-gods, rather than gods; creations of the great spirit of Ahura-
mazda. One power alone, the power of evil, seemed to contest his
supremacy. In opposition, therefore, to the god of that light which he
looked upon as the visible embodiment of the Good, the Persian
conceived the existence of a god of darkness, a god of evil, a god of
the under-world, a god of death. This god, Angro-mainyus,
possessed, indeed, the attributes both of Satan and of Pluto. There
was no hope for the complete triumph of good over evil in this life.
“Choose ye this day whom ye will serve,”—in that lay the whole
alternative, the ultimate possibility for good or evil, in so far as the
world of the present was concerned. Only in an after-life could the
final triumph of the good be looked for,—in a life after that
resurrection of the dead which the prophets, the sons of Zoroaster,
awaking from their long sleep, should bring to pass.
It seems at first strange that the Persian
POLYKRATES OF
SAMOS.
creed never captured the imagination of the
Greek. It may, indeed, be doubted whether it
was ever presented to him in its highest and purest form. The ideal
was, perhaps, too elevated for the ordinary devotee, and its full
appreciation confined to the initiated few. The Greek learnt indeed in
after times to admire certain of the virtues which the Persian
displayed; but never grasped, apparently, the spiritual and
intellectual basis which underlay them. It was long, too, ere the bitter
hostility to the barbarian allowed the Greek to view him and his ways
with unprejudiced eyes; and in that lapse of time the barbarian had
deteriorated, and his life had become more and more tainted with the
baser side of Oriental civilization, which could only excite contempt
in the Hellenic mind.
Of the history of the Asiatic Greeks during the later years of Cyrus
and the brief reign of his successor, Cambyses, but little is known.
Samos alone, as has been said, retained its independence. During
the last years of Cyrus, somewhere about 533, a certain Polykrates
made himself tyrant of the island, and under his rule the Samians
enjoyed a short period of prosperity, so great that it remained
proverbial in after-history. Polykrates used to the full the opportunity
afforded him by Cyrus’ detention in the East. Separated as he was
by only a few miles of sea from the great empire, he could not but
recognize the danger of his position, a danger which was rendered
far greater by the fact that the acquisition of Phœnicia had given the
Persians that arm they had up to that time lacked, a fleet. The great
prosperity of the island, due, no doubt, in a great measure to its
being the only Greek trading community on the Asiatic side which
was not under the Persian dominion, enabled him to raise and
maintain a large body of mercenaries as well as a fleet of a hundred
fifty-oared war-ships. He furthermore entered into negotiations with
Egypt, with a view to mutual defence.
At home in the Ægean, he played a many-sided part. Piracy,
trade, engineering, and territorial acquisition were all included in the
field of his manifold activity. The piracy was probably carried on at
the expense of those traders who did not use Samos as an entrepôt
between East and West. It involved him in many a quarrel with the
Asiatic Greek towns, whose anomalous position at the time is shown
by the recorded fact that Polykrates actually took possession of parts
of their territory on the mainland, although they were under the
Persian dominion. This somewhat wild career was interrupted, if not
positively checked, by events which were preparing on the far side of
the Levant.
Cambyses had made up his mind to complete by the conquest of
Egypt the work which his father had done. With a view to so doing,
he was collecting a great armament, in which a powerful fleet was to
play a part. This method of invasion, thus adopted for the first time,
served as a precedent for all the great Persian expeditions of after
years. The fact that it was Cambyses who conceived the design is
sufficient to stamp the picture which Herodotus draws of him as a
copy of a somewhat clumsy Egyptian caricature, even if other
evidence did not tend the same way.
Ships were levied from Phœnicia, and the Greeks of Cyprus had
H. iii. 19.
also to contribute to the fleet. The latter, after the fall of
Assyria, to which they had been in a position of
nominal subordination, had enjoyed a short period of absolute liberty.
Amasis of Egypt had reduced them to subjection; but, on the
establishment of the Persian power in the Syrian region, they had
thrown off their allegiance to Egypt and tendered their submission to
the new empire.
Polykrates began to reconsider his position. A Greek legend,
which Herodotus has preserved, represents him as having been
thrown over by Amasis out of superstitious apprehension. His
hitherto unvarying success, so thought the Egyptian king, must end
in some terrible disaster proportionate in greatness. Herodotus could
not resist a tale which so entirely harmonized with his views of life.
In actual fact, the reverse seems to have been the case.
Polykrates broke off the alliance with Amasis;
SAMOS ANNEXED
and not merely did so, but actually despatched
BY PERSIA.
forty ships to aid the Persian expedition. He
tried, indeed, to kill two birds with one stone, and missed both; for he
manned these vessels with Samian suspects, who had no mind to
lend their bodies for this experiment in diplomacy, and forthwith
turned Polykrates’ own weapons against himself by sailing back with
the fleet and making an attack on Samos. Failing in that, they sailed
away to Laconia, with a view to getting help of Sparta. Polykrates’
great bid for Persian favour had miscarried.
What followed is peculiarly interesting as being the first example
of the way in which Corinth could, and did, force the hand of the
Lacedæmonians in matters of policy.
The Lacedæmonians had indeed grievances against the
Samians; but it is unlikely that they would have undertaken the
expedition, had they not been urged thereto by Corinth. The
grievance on the side of Corinth was of a kind that was fated to
reappear on many momentous occasions in the course of the next
century. Corinthian trade had been interfered with by the Samians.
The piratical enterprise of Polykrates was sure to be directed against
the trade of a state which had broken off its old commercial relations
with Samos and transferred its connection to Miletus.
The expedition took place about 524. It failed. After a fruitless
siege of forty days the Lacedæmonians returned to Peloponnese.
Soon afterwards Polykrates met his end. He was enticed to
Magnesia on the Mæander by Orœtes, Satrap of Sardes, and there
put to death.
His secretary, Mæandrios, carried on the tyranny for some years;
but about the year 516 a Persian force invaded the island, and
H. iii. 139–
established Syloson, a brother of Polykrates who had
149. won the favour of Darius, as tyrant in the Persian
interest. A brother of Mæandrios made one vigorous
but vain attempt to win the island back. The acquisition of Samos
completed the Persian conquest of the Asian coast.
It is significant that Sparta, when appealed to by Mæandrios for
help, not merely refused it, but dismissed him from the Peloponnese,
lest he should bring about political complications. Sparta’s policy on
this occasion, and her attitude at the time of the Ionian revolt, show
that the fear of experience had taken the place with her of that
courage of ignorance which she had shown in her alliance with
Lydia.
Cambyses’ short reign came to an end in 522. He had added to
his dominions Egypt and the Libyan coast as far as the Greater
Syrtis, and had even made an expedition up the Nile into Ethiopia.
The Greeks in Egypt had been involved in the disaster which fell
upon their adopted home.
On the other hand, Persian enterprise in the West was for the
time being at a standstill; and the Asiatic Greeks, with the exception
of the Samians, seem to have passed seven uneventful years of
submission to their new ruler.
The last few months of Cambyses’ life had been troubled by the
plots of a Magian named Gaumata, who is said to have borne a
remarkable resemblance to Smerdis, a brother whom Cambyses had
caused to be murdered. The rising was no doubt encouraged by the
state of Cambyses’ health. He had certainly suffered from serious
illness in Egypt; there is, indeed, reason to suspect that he was an
epileptic.
The story of this false Smerdis is one of the unsolved mysteries of
the period. But few reliable details of it survive, and these are for the
most part contained in the great inscription of Darius on the rocks at
Behistun.
The usurpation was certainly popular in the home provinces of the
empire, if the rapid spread of the insurrection be any criterion. On his
way home from Egypt to suppress it, Cambyses died,—a violent
death, it seems certain, though historians are not in agreement as to
its exact form.
For some years the pseudo-Smerdis concealed his identity, and
maintained his power; but at last the suspected deception was
discovered by some of the great Persian nobles, among whom was
Darius, who claimed descent from the Achæmenid family. Of the
events that followed, nothing is known for
DARIUS.
certain, save that these nobles assassinated
the pretender, and Darius succeeded to the supreme power.
He seems to have been at first a king without a kingdom; for the
great satraps of the provinces, whose position placed at their
disposal large resources of men and money, revolted with well-nigh
one consent. The province in Asia Minor was one of the few which
H. iii. 126,
did not join in the rising. If a tale preserved in
128. Herodotus be true, its governor Orœtes meditated
insurrection; but before he could carry his plans into
action, he was assassinated by his own bodyguard, in obedience to
written orders sent by Darius.
The whole of the work of Cyrus seemed undone. The conquest of
the great Empire had to be carried out again, as it were, from the
beginning. How Darius carried it out is no real part of the present
story. Suffice it to say that he did the work, and that he seems to
have done it thoroughly.
This formidable upheaval showed Darius the necessity of giving
the empire a new organization, under which its recurrence would be
difficult or impossible. The time of Cyrus had been fully taken up with
the military acquisition and maintenance of the great realm.
Cambyses had been similarly occupied during his short reign;
though it may also be doubted whether he possessed the capacity
required for carrying out so huge a scheme. Under these two rulers
the old Assyrian method, or want of method, of administration had
largely prevailed, a system which seems to have been admirably
designed for goading the subject populations into rebellion, but
which provided no machinery by which insurrection could be
rendered difficult or be nipped in the bud. The central power was
ever kept on the strain by repeated revolts in the provinces, if the
term “province” can be applied to regions which were not in any real
sense “areas of administration,” but were merely regarded as lands
to be exploited for the benefit of the conquerors.
There are two important reasons why, in dealing with the history
of the relations between Greek and Persian, prominence should be
given to this organization of the empire under Darius. It was, in the
first place, destined to be the permanent political system of the
Persian dominion for all the ages during which that empire continued
to be the neighbour of the Greek of Europe and of Asia. In the
second place, it is impossible to realize the ability of the Persian race
at its highest point of development, and the enlightened character of
some, at least, of its rulers, without fully appreciating the main details
of the great scheme of imperial and provincial government which
Darius promulgated. In certain respects, indeed, its methods may
seem rude when compared with those of later ages; but in judging of
them it must be borne in mind that it was designed for the
government of peoples most of whom recognized no law save that of
the strong hand, and furthermore, that its creators were creators in a
very literal sense of the term, in that their work was so far in advance
of anything on the same scale which had preceded it, that its
originality is beyond question. The decay of the empire for whose
government it was formed was not due to faults in the scheme itself,
but to the rapid deterioration of those who administered it. It erred
perhaps on the side of centralization; but then the East does not
understand, and never has understood, anything but centralization in
government. Still, this feature was, owing to special circumstances,
destined to prove fatal to it; for it was at the centre of the empire, in
the reigning family itself, that the decay set in which corrupted the
whole.
The first danger to be provided against by the new scheme was
caused by the isolation and comparative independence of the
provincial governors, especially in the remoter provinces of the
empire. It is a form of danger common to all great empires at all ages
of the world, one against which the central government must ever
make provision. Darius’ solution of the difficulty was conceived on
much the same lines as were followed in later days by the Cæsars in
dealing with the Cæsarean provinces of the Roman empire. Not
merely was the area of administration of the
ORGANIZATION OF
THE PERSIAN
governor limited within the province, but also
EMPIRE. his actions were placed under immediate
observation by the appointment of high
officials, with special departments of their own, who were not under
his control, but were directly responsible to the head of the empire.
The plan had its disadvantages from the point of view of the
governed; and there were doubtless many instances in which the
subjects of the Great King, like the Roman provincial of later days,
Tac Agric. 15.
complained that “discord and harmony between those
set over them were alike disastrous to those they
ruled.” For the end for which it was devised, however, the method
seems to have been effective.
The empire was divided into satrapies, whose number varied from
3
twenty to twenty-eight at different periods of Darius’ reign. Persia
proper was alone excluded, receiving special treatment as the home
of the ruling race.
The civil and military powers in these satrapies or provinces were
divided. Three independent officials, with separate departments of
administration, were appointed to each. In the case of important
provinces the satraps were generally drawn from great families
connected with the Achæmenids; but in the case of the others the
field of choice seems to have been practically unlimited, and
governors were selected from among the comparatively poor as well
as from the wealthy, from the subject races as well as from the
Persians. The instances in which persons not of Persian or Medic
extraction were appointed to these important posts seem, however,
to have been rare and exceptional. There was no set period for the
tenure of the office. The duration of the governor’s administration
depended on the pleasure of the Great King.
In all civil matters the governor had absolute authority. He
controlled the administration of the taxes and the dispensation of
justice; he possessed the power of life and death. As viceroy,
representing the king, he was allowed to maintain a court and
bodyguard, with other minor attributes of regal power. Under ordinary
circumstances he neither commanded nor even controlled the
military forces in his province; in fact, a common policy of the central
government seems to have been to place a personal enemy of the
governor in command of the troops, whose relations with the king
were, of course, direct. It may be that under exceptional
circumstances,—as, for instance, when Artaphernes was satrap of
Sardes at the time of the Ionian revolt,—the supreme direction of
military operations was entrusted to a governor of peculiarly high
distinction.
The secretary of state within the province was a third official who
had immediate relations to the king. Though nominally the chief
assistant of the governor, he was in reality appointed to watch his
conduct, and to report irregularities or suspicious circumstances to
the central government at Susa.

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(eBook PDF) Conservation Biology by

(eBook PDF) Conservation Biology by Bradley J.

(eBook PDF) Conservation Biology by Bradley J.

(eBook PDF) An Introduction to Conservation Biology by

(eBook PDF) Principles of Conservation Biology. 3nd

(eBook PDF) An Introduction to Conservation Biology 2nd

(eBook PDF) The Economy Today 15th by Bradley Schiller

(Original PDF) Clinical Experiences in Counseling by

(eBook PDF) Essentials of Economics 11th Edition by

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4 Global Patterns and Drivers of Biodiversity

Part II Importance of Biodiversity 115

5 The Many Values of Biodiversity

6 Biodiversity and Ecosystem Services 141

7 Ecological Economics 181

Cost-effectiveness analysis 186 Indirect use value 194

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Box 7.1 Case in Point Surrogate market techniques 203

Part III Threats to Biodiversity 213

Box 8.1 Conservation in Practice Empirical measures 233

9 Habitat Loss, Fragmentation, and Degradation 249

Box 9.1 Challenges & Opportunities Desertification 283

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Box 10.1 Challenges & Opportunities Theory of Sustainable Harvesting 322

11 Invasive Alien Species

12 Climate Change 369

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Part IV Approaches to Conservation 403

13 Species-Level Conservation 405

Environmental stochasticity 411 Models of Population Size 426

14 Community and Ecosystem Conservation 445

(Category I) 446 Marine protected areas 463

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15 Landscape-Scale Conservation 477

16 Ex Situ Conservation 521

17 Conservation and Sustainable Development 555

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Why did we write this book?

Who is this book for?

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What pedagogical approaches does the book use?

• Success Stories, included in every chapter, detail conservation biology’s most

In addition to the pedagogical approaches, students are provided with a companion

What is our hope for this book?

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Dr. Matthew S. Becker, Prof. Aaron M. Ellison, Heidi Marcum, PhD,

We are indebted to the team at Sinauer Associates/Oxford University Press that

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eBook (ISBN 978-1-60535-882-6)

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If we are to ultimately become one of longest-lived species in Earth’s history,

State of the Human Species

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Figure 1.1 Human dominance is but a small fraction of Earth's 10

World population growth (in billions)

4.56 4.0 3.5 3.0 2.5 2.0 1.5 1.0 0.5 0