(eBook PDF) Essentials of Ecology 4th Edition by

Michael Begon download

https://ebookluna.com/product/ebook-pdf-essentials-of-
ecology-4th-edition-by-michael-begon/

Download full version ebook from https://ebookluna.com

 Contents

Preface xi What do we mean by a ‘species’? 39

Acknowledgments xiii Allopatric speciation 40

Sympatric speciation? 44
2.4 The effects of climatic change
Part 1 Introduction 1 on the evolution and distribution
of species 46
Chapter 1 Ecology and how to do it 3 2.5 Continental drift, parallel and
1.1 What is ecology? 4 convergent evolution 49
1.2 Scales, diversity of approaches, 2.6 Conclusion 52
and rigor 7 Summary 52
Questions of scale 7 Review Questions 53
The diversity of ecological evidence 10
Statistics and scientific rigor 12
1.3 Ecology in practice 15
Part 2 Conditions and Resources 55

 uccessions on old fields in Minnesota:

S Chapter 3 Physical conditions and
a study in time and space 15 the availability of
 ubbard Brook: a long-term commitment to
H resources 57
study at the ecosystem scale 18
3.1 Environmental conditions 58
 anada’s Experimental Lakes Area: decades
C
What do we mean by ‘harsh,’ ‘benign,’
of exploring the consequences of human
and ‘extreme’? 58
activities on lakes 20
Effects of conditions 59
 n introduction of an exotic fish species
A
to New Zealand: investigation on multiple Conditions as stimuli 61
biotic scales 21 The effects of conditions on interactions
Why Asian vultures were heading between organisms 63
for extinction: The value of a Responses by sedentary organisms 64
modeling study 24 Animal responses to environmental
Summary 26 temperature 64
Review questions 27 Microorganisms in extreme
environments 68
3.2 Resources for photosynthetic
Chapter 2 Ecology’s evolutionary organisms 68
backdrop 28 Solar radiation 69

2.1 Evolution by natural selection 29 Water 72

2.2 Evolution within species 32 Nutrients 75

Geographical variation within Carbon dioxide 75

species 32 3.3 Heterotrophs and their resources 80
Variation within a species with ­man-made Nutritional needs and provisions 80
selection pressures 37 Defense 83
Evolution and coevolution 39 3.4 Effects of intraspecific competition for
2.3 The ecology of speciation 39 resources 86

vi
 Contents  ■ vii  

3.5 Conditions, resources, and the ecological 5.1 Populations, individuals, births
niche 88 and deaths 124
Summary 89 What is an individual? 126
Review questions 89 Counting individuals, births, and deaths 126
5.2 Life cycles 128
Life cycles and reproduction 128
Chapter 4 Climate and the world’s
Annual life cycles 129
biomes 91
Longer life cycles 130
4.1 The world’s climate 92
5.3 Monitoring birth and death: life tables and
Redistribution of heat through
fecundity schedules 133
atmospheric movement 93
Cohort life tables 134
Ocean currents and the redistribution
Life tables for populations with
of heat 95
overlapping generations 137
4.2 Terrestrial biomes 95
A classification of survivorship curves 138
Biomes and convergent evolution 99
5.4 Dispersal and migration 138
Tropical rain forest 99
Dispersal determining abundance 140
Savanna 104
The role of migration 142
Temperate grasslands 105
5.5 The impact of intraspecific competition on
Desert 105
populations 142
Temperate forest 106
Patterns of population growth 143
Boreal forest (taiga) 106
5.6 Life history patterns 147
Tundra 107
Summary 152
The future distribution of terrestrial
Review Questions 152
biomes 108
4.3 Aquatic ecosystems on the
Chapter 6 Interspecific competition 154
continents 108
Streams and rivers 108 6.1 Ecological effects of interspecific
competition 155
Lakes and ponds 112
Competition amongst phytoplankton
Wetlands 113
for phosphorus 155
4.4 Ocean biomes 113
Coexistence and exclusion of competing
The deep ocean 114 salmonid fishes 156
Subtropical gyres 115 Some general observations 157
Coastal upwelling systems 116 Coexistence of competing diatoms 158
Broad continental shelves 117 Coexistence of competing birds 159
Nearshore coastal marine Competition between unrelated
ecosystems 117 species 159
Summary 119 The competitive exclusion principle 160
Review questions 120 Environmental heterogeneity 165
6.2 Evolutionary effects of interspecific
Part 3 Individuals and competition 169

Populations 121 Character displacement and ecological

release in the Indian mongoose 169
Chapter 5 Birth, death, and Character displacement in Canadian
movement 123 sticklebacks 170
viii ■ Contents

Evolution in action: selection on Chapter 8 Molecular and evolutionary

microorganisms 170 ecology 216
6.3 Interspecific competition and community
8.1 Molecular ecology: ­differentiation within
structure 172
and between species 217
Limiting resources and the regulation of
Differentiation within species:
diversity in phytoplankton
albatrosses 221
communities 172
Differentiation between species: the red
Niche complementarity amongst anem-
wolf—species or hybrid? 223
one fish in Papua New Guinea 172
8.2 Coevolutionary arms races 226
Species separated in space or in time 174
Coevolution 226
Spatial separation in trees and tree-root
fungi 175 Insect–plant arms races 226
Temporal separation in mantids and Coevolution of parasites and their
tundra plants 175 hosts 229
6.4 How significant is interspecific competition 8.3 Mutualistic interactions 232
in practice? 176 Mutualistic protectors 232
The prevalence of current competition 177 Farming crops or livestock 233
Competition or mere chance? 180 The dispersal of seeds and
Summary 183 pollen 235
Review questions 184 Mutualistic gut inhabitants 236
Mycorrhizas 236
Fixation of atmospheric nitrogen in
Chapter 7 Predation, grazing, and mutualistic plants 237
disease 185 Summary 240
7.1 What do we mean by predation? 186 Review questions 240
7.2 Prey fitness and abundance 187
7.3 The subtleties of predation 189
Part 4 Communities and
Interactions with other factors 190
Ecosystems 243
Compensation and defense by
individual prey 191 Chapter 9 From populations to
From individual prey to prey ­communities 245
populations 192
9.1 Multiple determinants of the dynamics of
7.4 Predator behavior: foraging and populations 246
transmission 195
Fluctuation or stability? 247
Foraging behavior 197
Determination and regulation of
7.5 Population dynamics of predation 199 abundance 248
Underlying dynamics of predator–prey Using k-value analysis 250
interactions: a tendency to cycle 199
9.2 Dispersal, patches, and metapopulation
Predator–prey cycles in dynamics 255
practice 203
9.3 Temporal patterns in community
Disease dynamics and cycles 204 composition 259
Crowding 207 Founder-controlled and
Predators and prey in patches 209 dominance-controlled
7.6 Predation and community structure 211 communities 259
Summary 214 Community succession 262
Review questions 215 9.4 Food webs 269
 Contents  ■ ix  

Indirect and direct effects 269 11.7 The flux of matter through ecosystems 332
Population and community stability and 11.8 Nutrient budgets and cycling at the
food web structure 274 ecosystem scale 334
Summary 280 Summary 338
Review questions 281 Review questions 339

Chapter 10 Patterns in species Part 5 Applied Issues in Ecology 341

richness 282 Chapter 12 Global biogeochemical

10.1 Quantifying species richness and cycles and their alteration by
diversity 283 humans 343
10.2 Spatially varying factors influencing species
12.1 What is biogeochemistry? 344
richness 285
12.2 The global carbon dioxide cycle 345
Productivity and resource richness 286
Understanding the carbon dioxide
Energy 288
sinks 349
Predation intensity 292
Effects of ocean acidification 351
Spatial heterogeneity 294
Will the terrestrial carbon dioxide sink
Environmental harshness 295 change in the future? 351
10.3 Temporally varying factors influencing Can we reduce carbon dioxide
species richness 296 emissions? 354
Climatic variation and its absence 296 12.3 The global methane cycle 354
Disturbance 296 The natural sources of methane 356
Environmental age: evolutionary time 297 Anthropogenic sources of methane 357
10.4 Habitat area and ­remoteness: Island Methane and the global climate
biogeography 298 system 357
10.5 Gradients of species richness 303 How do we reduce methane
Latitudinal gradients 303 emissions? 360
Gradients with altitude and depth 304 12.4 The nitrogen cycle at global
Gradients during community and regional scales 361
succession 306 Human acceleration of the nitrogen
Summary 307 cycle 362

Review questions 308 The ecological and human-health

costs of nitrogen 364
Regional variation in nitrogen pollution 365
Chapter 11 The flux of energy and matter How can we reduce nitrogen
through ecosystems 309 pollution? 367

11.1 The role of energy in ecology 310 Summary 368

11.2 Geographic patterns in ­primary Review questions 368

productivity 311
11.3 Factors limiting ­terrestrial primary
Chapter 13 Conservation ecology 370
productivity 312
11.4 Factors limiting aquatic 13.1 The need for conservation 371
primary productivity 318 13.2 Small populations 375
11.5 The fate of primary ­productivity: The classification of risk 375
grazing 325 Demographic risks associated with
11.6 The process of decomposition 330 small populations 375
x ■ Contents

Genetic problems in small populations 376 Two future inevitabilities 411

Habitat reduction 378 A global carrying capacity? 414
13.3 Threats to biodiversity 378 14.3 Ecology and human health 414
Overexploitation 378 Loss of the ozone layer 414
Habitat disruption 379 Extreme events 415
Global environmental change 382 Changing global patterns of infection 416
Introduced and invasive species 383 Emerging infectious diseases 418
Infectious disease 386 14.4 Synthetic fertilizer and the intensification
Combinations of risks and extinction of agriculture 418
vortices 386 14.5 Monocultures, pests, and pesticides in
Chains of extinctions? 386 agriculture 421

13.4 Conservation in practice 389 Chemical approaches to pest

control 422
Population viability analysis 390
Biological control 426
Protected areas 393
14.6 Global land use and other constraints
Selecting conservation areas 397
on ­continued intensification of
Collections of areas 400 agriculture 428
13.5 Ecosystem services 401 14.7 Food from fisheries and aquaculture 434
Summary 404 Summary 439
Review questions 404 Review questions 440

Chapter 14 The ecology of human

­population growth, disease, References R-1
and food supply 406 Index I-1
14.1 Human use of ecological resources 407
14.2 The human population problem 408 Glossary G-1
Population growth up to the present 408 (available online only: www.wiley.com/
Predicting the future 410 college/begon)
 Preface

By writing this book we hope to share with you some counter and solve these problems depend absolutely on
of our wonder at the complexity of nature, but we must a proper grasp of ecological fundamentals.
all also be aware that there is a darker side: the fear The book is divided into five sections. In the
that we are destroying our natural environments and introduction we deal with two foundations for the sub-
the services they provide. All of us need to be ecologi- ject that are often neglected in texts. Chapter 1 aims to
cally literate so that we can take part in political debate show not only what ecology is but also how ecologists
and contribute to solving the ecological problems that do it—how ecological understanding is achieved, what
we carry with us in this new millennium. We hope our we understand (and, just as important, what we do not
book will contribute to this objective. yet understand) and how our understanding helps us
The genesis of this book can be found in the predict and manage. We then introduce ‘Ecology’s evo-
more comprehensive treatment of ecology in our big lutionary backdrop’ and show that ecologists need a
book Ecology: from Individuals to Ecosystems (Begon, full understanding of the evolutionary biologist’s disci-
Townsend & Harper, 4th edn, 2006). This is used as an pline in order to make sense of patterns and processes
advanced university text around the world, but many of in nature (Chapter 2).
our colleagues have called for a more succinct treatment What makes an environment habitable for par-
of the essence of the subject. Thus, we were spurred into ticular species is that they can tolerate the physico-
action to produce a distinctively different book, writ- chemical conditions there and find in it their essential
ten with clear objectives for a different audience—those resources. In the second section we deal with condi-
taking a semester-long beginners course in the essentials tions and resources, both as they influence individual
of ecology. We hope that at least some readers will be species (Chapter 3) and in terms of their consequences
excited enough to go on to sample the big book and the for the composition and distribution of multispecies
rich literature of ecology that it can lead into. communities and ecosystems, for example in deserts,
In this fourth edition of Essentials of Ecology rain forests, rivers, lakes and oceans (Chapter 4).
we have continued to make the text, including math- The third section (Chapters 5–8) deals system-
ematical topics, accessible while updating the material atically with the ecology of individual organisms
and expanding our coverage of ecosystem science and and populations, with chapters on ‘birth, death and
biogeochemistry. The fourth edition extensively cov- movement’ (Chapter 5), ‘interspecific competition’
ers both terrestrial and aquatic ecology, and we have (Chapter 6), and ‘predation, grazing, and disease’
strived to demonstrate how ecological principles apply (Chapter 7). This section also includes a chapter on
equally to both types of environments. While we have ‘Molecular and evolutionary ecology’, added origi-
expanded coverage on some topic areas in the fourth nally in the third edition and responding to the feelings
edition, we worked hard to not expand the size of the of some readers that, although evolutionary ideas per-
book. We want this text to be a readily accessible read. vade the book, there was still not sufficient evolution
Ecology is a vibrant subject and this is reflected for a book at this level.
by our inclusion of literally hundreds of new studies. In the fourth section (Chapters 9–11), we move
Some readers will be engaged most by the fundamen- up the hierarchical scale of ecology to consider commu-
tal principles of how ecological systems work. Others nities consisting of many populations, and ecosystems,
will be impatient to focus on the ecological problems where we focus on the fluxes of energy and matter
caused by human activities. We place heavy emphasis between and within systems.
on both fundamental and applied aspects of ecology: Finally, armed with knowledge and understand-
there is no clear boundary between the two. However, ing of the fundamentals, the book turns to the appli-
we have chosen to deal first in a systematic way with cation of ecological science to some of the major
the fundamental side of the subject, and we have done environmental challenges of our time. Our goal in
this for a particular reason. An understanding of the these final chapters is not to provide encyclopedic
scope of the problems facing us (the unsustainable coverage to these environmental problems, but rather
use of ecological resources, pollution, extinctions and to illustrate how ecology contributes to understand-
the erosion of natural biodiversity) and the means to ing the problems, and can potentially help with their

xi
xii ■ Preface

solution. In Chapter 12, we focus on global biogeo- • You will also find three categories of boxed text:
chemical cycles, such as the global carbon dioxide
cycle and how this has been dramatically changed • ‘Historical landmarks’ boxes emphasize some
by burning fossil fuels and other human activities. landmarks in the development of ecology.
In ‘conservation ecology’ (Chapter 13), we develop • ‘Quantitative aspects’ boxes set aside mathemati-
an armory of approaches that may help us to save cal and quantitative aspects of ecology so they do
endangered species from extinction and conserve not unduly interfere with the flow of the text and
some of the biodiversity of nature for our descendants. so you can consider them at leisure.
The final chapter, ‘the ecology of human population
growth, disease, and food supply,’ takes an ecological • ‘ECOncerns’ boxes highlight some of the applied
approach to examining the issues of the population problems in ecology, particularly those where there
problem, of human health, and of the sustainability of is a social or political dimension (as there often is). In
agriculture and fisheries. these, you will be challenged to consider some ethical
A number of pedagogical features have been questions related to the knowledge you are gaining.
included to help you. An important further feature of the book is the
• Each chapter begins with a set of key concepts that companion web site, accessed through Wiley at www
you should understand before proceeding to the .wiley.com/college/begon. This provides an easy-to-use
next chapter. range of resources to aid study and enhance the content of
the book. Features include self-assessment multiple choice
• Marginal headings provide signposts of where you questions for each chapter in the book, an interactive
are on your journey through each chapter—these
tutorial to help students to understand the use of math-
will also be useful revision aids.
ematical modeling in ecology, and high-quality images of
• Each chapter concludes with a summary and a set the figures in the book that teachers can use in preparing
of review questions, some of which are designated their lectures or lessons, as well as access to a Glossary
challenge questions. of terms for use with this book and for ecology generally.
 Acknowledgments

It is a pleasure to record our gratitude to the people strong, and we gratefully acknowledge his tremendous
who helped with the planning and writing of this contribution to the series.
book. Going back to the first edition, we thank Bob We are also grateful to the following colleagues
Campbell and Simon Rallison for getting the origi- who provided insightful reviews of early drafts of
nal enterprise off the ground and Nancy Whilton one or more chapters in this or earlier editions, or
and Irene Herlihy for ably managing the project; and who gave us important advice and leads: William
for the second edition, Nathan Brown (Blackwell, Ambrose (Bates College), Vickie Backus (Middlebury
US) and Rosie Hayden (Blackwell, UK) for making College), James Cahill (University of Alberta), Liane
it so easy for us to take this book from manuscript Cochrane-Stafira (Saint Xavier University), Mark Davis
into print. For the third edition, we especially thank (Macalester College), Tim Crews (The Land Institute),
Nancy Whilton and Elizabeth Frank in Boston for Kevin Dixon (Arizona State University, West), Stephen
persuading us to pick up our pens again (not liter- Ellner (Cornell University), Alex Flecker (Cornell
ally) and Rosie Hayden, again, and Jane Andrew and University), Bruce Grant (Widener University), Christy
Ward Cooper for seeing us through production. For Goodale (Cornell University), Don Hall (Michigan
this fourth edition, we thank Rachel Falk (Wiley, State University), Jenny Hodgson, Greg Hurst (both
USA) for getting the ball rolling and for bringing University of Liverpool), William Kirk (Keele University,
in one of us (RWH) as a new author, Elisa Adams UK), Hans deKroon (University of Nijmegen), Zen
for her superb assistance with text editing, Chloe Lewis (University of Liverpool), Sara Lindsay (Scripps
Moffett, Elizabeth Baird, MaryAnn Price and Lisa Institute of Oceanography), James Maki (Marquette
Torri (Precision Graphics) for their excellent over- University), George Middendorf (Howard Univer­
seeing of the final production, and the entire Wiley sity), Paul Mitchell (Staffordshire University, UK),
team for their dedicated efforts and cheerful “can- Tim Mousseau (University of South Carolina), Katie
do” attitude. O’Reilly (University of Portland), Clayton Penniman
We note with sadness the passing in 2009 of (Central Connecticut State University), Tom Price
our long-time mentor and collaborator John Harper, (Univeristy of Liverpool), Jed Sparks (Cornell Univer­
author on the first three editions of this book. We owe sity), Catherine Toft (UC Davis), David Tonkyn
him a special debt of gratitude that extends far beyond (Clemson University), Saran Twombly (University of
the past co-authorship of this book into all aspects of Rhode Island), Jake Weltzin (University of Tennessee at
our lives as ecologists. He is sorely missed. Knoxville), and Alan Wilmot (University of Derby, UK).
Colin Townsend, the lead author on the first three Last, and perhaps most, we are glad to thank our
editions of Essentials of Ecology, has stepped from wives and families for continuing to support us, listen
the treadmill of revisions and let us take the lead on to us, and ignore us, precisely as required—thanks to
this fourth edition. His imprint on the book remains Linda, and to Roxanne and Marina.

Michael Begon, Liverpool, UK and

Robert Howarth, Ithaca, NY USA

xiii
 1
2
Ecology and how to do it 3
Ecology’s evolutionary
backdrop 28 Part 1
Introduction
Conditions and Resources
© ElementalImaging/iStockphoto
 © ElementalImaging/iStockphoto

Chapter 1
Ecology and how to do it
CHAPTER CONTENTS
1.1 What is ecology? 1.3 Ecology in practice
1.2 Scales, diversity of approaches, and rigor

KEY CONCEPTS
After reading this chapter you will be able to:

• explain how ecologists seek to describe and • describe how ecologists use observations, field
understand, and on the basis of their under- and laboratory experiments, and mathematical
standing, to predict, manage, mitigate, and models to collect scientific evidence
control

• describe the variety of spatial and temporal

scales on which ecological phenomena occur
4 ■ Part 1 Introduction

E
cology today is a subject about which almost everyone has heard and most
people consider to be important—even when they are unsure about the
exact meaning of the term. There can be no doubt that it is important, but this
makes it all the more critical that we understand what ecology is and how to do it.

1 .1 WHAT I S E CO LO G Y ? are good at preparing you for an examination, they are

not so good at capturing the flavor and excitement of
We could answer the question ‘What is ecology?’ by ecology. There is a lot to be gained by replacing that
examining various definitions that single question about a definition with a series of more
the earliest
have been proposed and choosing ecologists provocative ones: ‘What do ecologists do?’ ‘What are
one as the best (Box 1.1). But while ecologists interested in?’ and ‘Where did ecology
definitions have conciseness and precision, and they emerge from in the first place?’

1.1 Historical Landmarks

Definitions of ecology

Ecology (originally in German, Öekologie) was first defined in 1866 by Ernst Haeckel, an enthusiastic and
influential disciple of Charles Darwin. To him, ecology was ‘the comprehensive science of the relationship
of the organism to the environment.’ The spirit of this definition is very clear in an early discussion of bio-
logical subdisciplines by Burdon-Sanderson (1893), in which ecology is ‘the science which concerns itself
with the external relations of plants and animals to each other and to the past and present conditions of
their existence,’ to be contrasted with physiology (internal relations) and morphology (structure).
In the years after Haeckel, plant ecology and animal ecology drifted apart. Influential works defined
ecology as ‘those relations of plants, with their surroundings and with one another, which depend directly
upon differences of habitat among plants’ (Tansley, 1904), or as the science ‘chiefly concerned with what
may be called the sociology and economics of animals, rather than with the structural and other adapta-
tions possessed by them’ (Elton, 1927). The plant ecologists and animal ecologist, though, have long since
agreed that they belong together, and more recent definitions of ecology include all organisms, including
bacteria, archaea, algae, and fungi in addition to plants and animals. Most modern definitions stress the
relationships between and among organisms. For example, two textbooks from the 1970s defined ecology
as ‘the study of the natural environment, particularly the interrelationships between organisms and their
surroundings’ (Ricklefs, 1973) and as ‘the scientific study of the interactions that determine the distribution
and abundance of organisms’ (Krebs, 1972).
Ecology certainly includes the investigation of organisms and their interactions, but to many ecolo-
gists, definitions that focus only on these interactions and on the distribution and abundance of organisms
are too narrow. Ecologists also examine the interaction between life and the physical environment, for
instance studying how organisms affect material fluxes in nature. The sequestration of carbon dioxide by a
forest would be one example of this. Beginning in the mid-20th century, the American ecologist E. P. Odum
(1953) pushed for a broader definition of ecology:‘the study of the structure and function of nature, which
 Ecology and how to do it Chapter 1  ■ 5  

includes the living world.’ Many have thought this definition overly broad, as geologists and meteorologists
also study aspects of the structure and function of nature. In 1992, G. E. Likens stressed the need for the
definition of ecology to include ‘the interactions between organisms and the transformation and flux of
energy and matter.’ We agree, and in this text define ecology as:

the scientific study of the distribution and abundance of organisms, the interactions that determine
that distribution and abundance, and the relationships between organisms and the transformation
and flux of energy and matter.

Ecology can lay claim to being the oldest science, preparation. There is nothing . . . more erroneous
as the most primitive humans must have been ecolo- than this feeling.
gists of sorts, driven by the need to understand where
and when their food and their (nonhuman) enemies On the other hand, the need for applied ecology
were to be found. The earliest agriculturalists needed to to be based on its pure counterpart was clear in the
be even more sophisticated, with knowledge of how introduction to Charles Elton’s (1927) Animal Ecology
to manage their domesticated sources of food. These (Figure 1.1):
early ecologists, then, were applied ecologists, seeking Ecology is destined for a great future . . . The
to understand the distribution, abundance, and produc- tropical entomologist or mycologist or weed-
tivity of organisms in order to apply that knowledge for controller will only be fulfilling his functions
their own benefit. Applied ecologists today still have properly if he is first and foremost an ecologist.
many of the same interests: how to optimize the rate at
which food is collected from natural environments in a In the intervening years, the coexistence of these
sustainable way; how domesticated plants and animals pure and applied threads has been maintained and
can best be managed so as to maximize rates of return; built upon. Many applied sciences such as forestry,
how food organisms can be protected from their own agronomy, and fisheries biology have contributed to
natural enemies; and how to control the populations of the development of ecology and have seen their own
pathogens and parasites that live on us.
In the last century or so, how-
a pure and applied
ever, since ecologists have been science
self-conscious enough to give them-
selves a name, ecology has consistently covered not
only applied but also fundamental, ‘pure’ science.
A.G. Tansley was one of the founding fathers of ecology.
He was concerned especially to understand, for under-
standing’s sake, the processes responsible for determin-
ing the structure and composition of different plant
communities. When, in 1904, he wrote from Britain
about ‘The problems of ecology’ he was particularly
worried by a tendency for too much ecology to remain
at the descriptive and unsystematic stage (such as accu-
mulating descriptions of communities without know-
ing whether they were typical, temporary, or whatever),
courtesy Robert Elton

too rarely moving on to experimental or systematically

planned, or what we might call a scientific analysis.
Tansley’s worries were echoed in the United States
by another of ecology’s founders, F. E. Clements, who in
1905 in his Research Methods in Ecology complained:
F I GURE 1. 1 One of the great founders of ecology: Charles
The bane of the recent development popularly Elton (1900–1991). Animal Ecology (1927) was his first book but
known as ecology has been a widespread feeling The Ecology of Invasions by Animals and Plants (1958) was equally influ-
that anyone can do ecological work, regardless of ential. (After Breznak, 1975.)
6 ■ Part 1 Introduction

development enhanced by ecological ideas and app­ parasites and predators that attack it. This is a proxi-
roaches. All aspects of food and fiber gathering, pro- mate explanation – an explanation in terms of what
duction, and protection have been involved. The is going on ‘here and now.’ We can also ask how this
biological control of pests (the use of pests’ natural bird came to have these properties that now govern
enemies to control them) has a history going back at its life. This question has to be answered by an expla-
least to the ancient Chinese but has seen a resurgence nation in evolutionary terms; the ultimate explanation
of ecological interest since the shortcomings of chemi- of the present distribution and abundance of this bird
cal pesticides began to be widely apparent in the 1950s. lies in the ecological experiences of its ancestors (see
The ecology of pollution has been a growing concern Chapter 2).
from around the same time and expanded further in In order to understand something, of course,
the 1980s and 1990s from local to regional and global we must first have a description of whatever it is we
issues. The last few decades have also seen expansions wish to understand. Ecologists must therefore describe
in both public interest and ecological input into the before they explain. On the other hand, the most valu-
conservation of endangered species and the biodiver- able descriptions are those carried out with a particular
sity of whole areas, the control of disease in humans as problem or ‘need for understanding’ in mind. Undirected
well as many other species, and the potential conse- description, carried out merely for its own sake, is often
quences of profound human-caused changes to the later found to have selected the wrong things and has
global environment. little place in ecology—or any other science.
And yet, at the same time, Ecologists also often try to predict. For exam-
unanswered
many fundamental problems of questions ple, how will global warming affect the sequestra-
ecology remain unanswered. To tion (storage) of carbon in natural ecosystems? Will
what extent does competition for food determine warming reduce this storage, and therefore result in
which species can coexist in a habitat? What role does even more global warming since less carbon diox-
disease play in the dynamics of populations? Why are ide will be removed from the atmosphere? Often,
there more species in the tropics than at the poles? ecologists are interested in what will happen to a
What is the relationship between soil productivity and population of organisms under a particular set of cir-
plant community structure? Why are some species cumstances, and on the basis of these predictions to
more vulnerable to extinction than others? Are wet- control, exploit or conserve the population. We try
lands net sources or sinks of greenhouse gas emission to minimize the effects of locust plagues by predict-
to the atmosphere? And so on. Of course, unanswered ing when they are likely to occur and taking appro-
questions—if they are focused questions—are a symp- priate action. We try to exploit crops most effectively
tom of the health, not the weakness, of any science. But by predicting when conditions will be favorable to the
ecology is not an easy science, and it has particular crop and unfavorable to its enemies. We try to preserve
subtlety and complexity, in part because ecology is rare species by predicting the conservation policy
peculiarly confronted by ‘uniqueness’: millions of dif- that will enable us to do so. Some prediction and
ferent species, countless billions of genetically distinct control can be carried out without deep explanation
individuals, all living and interacting in a varied and or understanding: it is not difficult to predict that the
ever-changing world. The beauty of ecology is that it destruction of a woodland will eliminate woodland
challenges us to develop an understanding of very birds. But what if the woodland is not destroyed, but
basic and apparent problems—in a way that recog- rather fragmented into distinct parts with suburbs or
nizes the uniqueness and complexity of all aspects of agricultural fields between them? What effect may
nature – but seeks patterns and predictions within this this have on the woodland birds? Insightful predic-
complexity rather than being swamped by it. tions, precise predictions, and predictions of what
Let’s come back to the ques- will happen in unusual circumstances can be made
understanding,
tion of what ecologists do. First description, only when we can also explain and understand what
and foremost ecology is a science, and ­prediction, and is going on.
ecologists therefore try to explain control This book is therefore about:
and understand. Explanation can
be either ‘proximate’ or ‘ultimate,’ and ecologists are 1 How ecological understanding is achieved.
interested in both. For example, the present distribu-
2 What we do understand, and what we do not.
tion and abundance of a particular species of bird may
be ‘explained’ in terms of the physical environment 3 How ecological understanding can help us predict,
that the bird tolerates, the food that it eats, and the manage, mitigate, and control.
 Ecology and how to do it Chapter 1  ■ 7  

1.2 SCA LE S , DI V E R S I T Y O F 1 The properties observed at a particular level arise

APP R OACH E S , AN D R I GOR out of the functioning of parts at the level below.
For example, how a tissue functions is the result of
Ecology is a diverse discipline, and ecologists use a vast the functioning of the cells in that tissue, and how
array or tools and approaches. Later in this chapter, we an ecosystem functions is the result of the function-
briefly give some examples of this diversity, but first ing of the communities within it interacting with the
we elaborate on three general points: physical environment.
• ecological phenomena occur at a variety of scales; 2 In order to understand the mechanistic reasons that
a particular property is observed at any level of bio-
• ecological evidence comes from a variety of differ-
ent sources; logical organization, a scientist needs to look at the
next lowest level of organization. To understand
• ecology relies on truly scientific evidence. dysfunction in an individual organism, we must
look at the functioning of the organs in that organ-
Questions of scale ism; and to understand the controls on birth rate
Ecology operates at a range of scales: time scales, spa- in a population, we must look at reproduction in
tial scales, and ‘biological’ scales. It is important to individual organisms.
appreciate the breadth of these and how they relate 3 However, properties observed at a given level of
to one another. organization may be predicted without fully under-
Life is studied at a variety of the ‘biological’ standing the functioning at lower levels. This third
hierarchical levels, with much of biol- scale generality may seem to contradict the other two, but
ogy focused on levels from molecules, it does not. Consider an analogy from the physical
to organelles, cells, tissues, organs, and whole organisms. sciences. As early as 1662, Boyle knew that when
Ecologists study levels from individual organisms, to the pressure of a gas is doubled, its volume is halved,
populations, communities, ecosystems, and the global if temperature remains constant. This behavior of
biosphere (Figure 1.2). the gas as a whole is the result of the interactions
• Populations are functioning groups of individual of the gas molecules, yet Boyle’s law provided valu-
organisms of the same species in a defined location. able predictive power for centuries, long before the
concept of the molecule was developed. Today,
• Communities consist of all the species populations physical chemists can indeed explain gas behavior
present in a defined location. based on understanding of the behavior of individ-
• Ecosystems include both the community of organisms ual molecules, but the explanation is complex, and
and the physical environment in which they exist. not even taught to most undergraduate college stu-
dents. Similarly, ecologists can predict patterns in
• The biosphere is the totality of all of life interacting ecosystems without understanding all of the details
with the physical environment at the scale of the
of the dynamics of constituent populations, and can
entire planet.
predict patterns in populations without understand-
At the level of the organism, ecology deals primarily ing all of the details of the responses of individual
with how individuals are affected by their environment organisms.
and with their physiological and behavioral responses
to the environment. Population ecology stresses the Within the living world, there a range of spatial
trends and fluctuations in the number of individual of is no arena too small nor one so scales
a particular species at a particular time and place, as large that it does not have an ecol-
determined by the interactions of birth and death rates ogy. Even the popular press talk increasingly about the
and the interactions between the populations them- ‘global ecosystem’, and there is no question that several
selves (such as predators and prey). Community ecology ecological problems can be examined only at this very
focuses on questions such as what controls the diversity large scale. These include the relationships between
of species of in a given area. Ecosystem ecology strives to ocean currents and fisheries, or between climate patterns
understand the functioning of entire lakes, forests, wet- and the distribution of deserts and tropical rain forests,
lands, or other portions of the Earth in terms of energy or between elevated carbon dioxide in the atmosphere
and material inputs and outputs. Across all scales of (from burning fossil fuels) and global climate change.
biological hierarchy—including these ecological ones— At the opposite extreme, an individual cell may
three generalities emerge. be the stage on which two populations of pathogens
 8

OTHER LEVELS OF BIOLOGY THE REALM OF ECOLOGY

■

NASA

FI GU R E 1.2
iy/Shutterstoc
o Yur k
ykayl
Br

e Source
cienc
n/S
sig

De
na
Lagu
Part 1 Introduction

ence Source
/Sci
ign
es

D
na
Lagu
pho
/iStock to
oto
ph
o_

cr
mi
Biosphere

old/
r Arn
/P
e Im
ete ages
Ecosystem

i
tty

Ecology is studied at many hierarchical levels.

dB
Ge rk
Rolan
Community

Source
nce
cie
s/S

e
iat
Population

soc
Biophoto As
Organism
ourc e
ce S
ien
Sc

/
ign
Laguna Des
Organs and tissues

Cells
ages
y Im
ett

c/G
lini
3DC
Organelles

Molecules
 Ecology and how to do it Chapter 1  ■ 9  

may be studied in butterflies over the course of days, or

in the forest trees that are still (slowly) migrating into
deglaciated areas following the last ice age.
The appropriate time scale the need for long-
for ecological investigation varies term studies
with the question to be answered.
However, many ecological studies end up being shorter
than appropriate for the question, due to human frail-
ties. Longer studies cost more and require greater
dedication and stamina. The often short-term nature
of funding, an impatient scientific community, and the
requirement for concrete evidence of activity for career
progression all put pressure on ecologists (and all sci-
Shigeharu Moriya

entists) to publish their work sooner rather than later.

Why are long-term studies potentially of such value?
The reduction over a few years in the numbers of a
particular species of wild flower, or bird, or butterfly
FI GU R E 1.3 The diverse community of a termite’s gut. Termites might be a cause for conservation concern—but one or
can break down lignin and cellulose from wood because of their mutu- more decades of study may be needed to be sure that the
alistic relationships (see Chapter 8) with a diversity of microbes that live decline is more than just an expression of the random
in their guts. ups and downs of ‘normal’ population dynamics. One
of the longest, continuously run ecological studies is
compete with one another for the resources that the at the Hubbard Brook Experiment Forest in the White
cell provides. At a slightly larger spatial scale, a ter- Mountains of New Hampshire. Among other mea-
mite’s gut is the habitat for bacteria, protozoans, and sures, Gene Liken and other scientists there have moni-
other species (Figure 1.3) – a community whose diver- tored the acidity of rain since the early 1960s. In the
sity is comparable to that of a tropical rain forest in 1960s, the rain was quite acidic (low pH: high hydro-
terms of the richness of organisms living there, the gen ion concentrations), and this was in fact one of
variety of interactions in which they take part, and the earliest discoveries anywhere of the phenomenon
indeed the extent to which we remain ignorant about of acid rain. The long-term trend, though, has been for
the species identity of many of the participants. precipitation to become less acidic over subsequent
Between these extremes, different ecologists, or the decades (Figure 1.4); but we can observe this only
same ecologist at different times, may study the inhab-
itants of pools that form in small tree-holes, the tempo-
We are able to observe a
rary watering holes of the savannas, or the great lakes long-term drop in rain acidity
100 only through studies over
H+ concentration in precipitation (µeq/L)

and oceans; others may examine the diversity of fleas many years.
on different species of birds, the diversity of birds in 90

different sized patches of woodland, or the diversity of 80

woodlands at different altitudes. 70
To some extent related to this
a range of time 60
range of spatial scales, and to the scales
levels in the biological hierarchy, 50

ecologists also work on a variety of time scales. 40

Ecological succession – the successive and continuous 30
colonization of a site by certain species populations,
20
accompanied by the local extinction of others – may be 1960 1965 1970 1975 1980 1985 1990 1995 2000 2005
studied over a period from the deposition of a lump of
F I GURE 1. 4 Hydrogen ion concentration in precipitation at the
sheep dung to its decomposition (a matter of weeks),
Hubbard Brook Experimental Forest over time. Note the long-term trend
from the abandonment of a patch of tropical rain for-
of decreasing concentration, indicating that the pH has been rising, and
est cleared for slash-and-burn agriculture (years to the rain has become less acidic over time. However, analysis of periods
decades), or from the development of a new forest on of only a few years in duration can show sharp increases or decreases
land wiped clean to bedrock by the retreat of a glacier in the hydrogen ion concentration, and are quite misleading with regard
in the arctic or high mountains (centuries). Migration to the long-term trend (After Likens 2004).
Discovering Diverse Content Through
Random Scribd Documents
True to his Colors 1 25
Rodney, the Partisan 1 25

Marcy, the Blockade Runner 1 25

Marcy, the Refugee 1 25

OUR FELLOWS; or, Skirmishes with the
Swamp Dragoons. By Harry Castlemon.
16mo. Fully illustrated. Cloth, extra 1 25
Specimen Cover of the Ragged
Dick Series.

Alger's
Renowned
Books.
 by

Horatio

Alger, Jr.

Horatio Alger, Jr., has attained distinction as one of the most

popular writers of books for boys, and the following list comprises all
of his best books.
⁂ Any volume sold separately.

RAGGED DICK SERIES. By Horatio Alger,

Jr. 6 vols., 12mo. Fully illustrated. Cloth,
extra, printed in colors. In box $7 50
Ragged Dick; or, Street Life in New York 1 25
Fame and Fortune; or, The Progress of
Richard Hunter 1 25

Mark, the Match Boy; or, Richard Hunter's

Ward 1 25

Rough and Ready; or, Life among the New

York Newsboys 1 25
Ben, the Luggage Boy; or, Among the
Wharves 1 25
Rufus and Rose; or, the Fortunes of Rough
and Ready 1 25
TATTERED TOM SERIES. (First Series.) By
Horatio Alger, Jr. 4 vols., 12mo. Fully
illustrated. Cloth, extra, printed in colors.
In box 5 00
Tattered Tom; or, The Story of a Street
Arab 1 25
Paul, the Peddler; or, The Adventures of a
Young Street Merchant 1 25

Phil, the Fiddler; or, The Young Street

Musician 1 25

Slow and Sure; or, From the Sidewalk to

the Shop 1 25
TATTERED TOM SERIES. (Second Series.) 4
vols., 12mo. Fully illustrated. Cloth, extra,
printed in colors. In box $5 00

Julius; or the Street Boy Out West 1 25

The Young Outlaw; or, Adrift in the World 1 25

Sam's Chance and How He Improved it 1 25

The Telegraph Boy 1 25
LUCK AND PLUCK SERIES. (First Series.)
By Horatio Alger, Jr. 4 vols., 12mo. Fully
illustrated. Cloth, extra, printed in colors.
In box $5 00
Luck and Pluck; or John Oakley's
Inheritance 1 25

Sink or Swim; or, Harry Raymond's Resolve 1 25

Strong and Steady; or, Paddle Your Own
Canoe 1 25
Strive and Succeed; or, The Progress of
Walter Conrad 1 25
LUCK AND PLUCK SERIES. (Second
Series.) By Horatio Alger, Jr. 3 vols.,
12mo. Fully illustrated. Cloth, extra,
printed in colors. In box $5 00
Try and Trust; or, The Story of a Bound
Boy 1 25
Bound to Rise; or Harry Walton's Motto 1 25

Risen from the Ranks; or, Harry Walton's

Success 1 25

Herbert Carter's Legacy; or, The

Inventor's Son 1 25

CAMPAIGN SERIES. By Horatio Alger, Jr. 3

vols., 12mo. Fully illustrated. Cloth, extra,
printed in colors. In box $3 75

Frank's Campaign; or, The Farm and the

Camp 1 25
Paul Prescott's Charge 1 25
Charlie Codman's Cruise 1 25

BRAVE AND BOLD SERIES. By Horatio $5 00

Alger, Jr. 4 vols., 12mo. Fully illustrated.
 Cloth, extra, printed in colors. In box

Brave and Bold; or, The Story of a Factory

Boy 1 25

Jack's Ward; or, The Boy Guardian 1 25

Shifting for Himself; or, Gilbert Greyson's
Fortunes 1 25
Wait and Hope; or, Ben Bradford's Motto 1 25

PACIFIC SERIES. By Horatio Alger, Jr. 4

vols. 12mo. Fully illustrated. Cloth, extra,
printed in colors. In box $5 00

The Young Adventurer; or, Tom's Trip

Across the Plains 1 25

The Young Miner; or, Tom Nelson in

California 1 25
The Young Explorer; or, Among the
Sierras 1 25
Ben's Nugget; or, A Boy's Search for
Fortune. A Story of the Pacific Coast 1 25
ATLANTIC SERIES. By Horatio Alger, Jr. 4
vols., 12mo. Fully illustrated. Cloth, extra,
printed in colors. In box $5 00
The Young Circus Rider; or, The Mystery
of Robert Rudd 1 25
Do and Dare; or, A Brave Boy's Fight for
Fortune 1 25

Hector's Inheritance; or, Boys of Smith 1 25

 Institute

Helping Himself; or, Grant Thornton's

Ambition 1 25
WAY TO SUCCESS SERIES. By Horatio
Alger, Jr. 4 vols., 12mo. Fully illustrated.
Cloth, extra, printed in colors. In box $5 00
Bob Burton 1 25
The Store Boy 1 25

Luke Walton 1 25
Struggling Upward 1 25

New Book by Alger.

DIGGING FOR GOLD. By Horatio Alger, Jr.

Illustrated 12mo. Cloth, black, red and
gold 1 25
Specimen Cover of the Wyoming
Series.

A
New Series
of Books.

Indian Life
and
Character
 Founded on
Historical
Facts.

By Edward S. Ellis.
⁂ Any volume sold separately.

BOY PIONEER SERIES. By Edward S. Ellis.

3 vols., 12mo. Fully illustrated. Cloth,
extra, printed in colors. In box $3 75

Ned in the Block House; or, Life on the

Frontier. 1 25

Ned in the Woods. A Tale of the Early

Days in the West 1 25
Ned on the River 1 25
DEERFOOT SERIES. By Edward S. Ellis. In
box containing the following. 3 vols.,
12mo. Illustrated $3 75
Hunters of the Ozark 1 25

Camp in the Mountains 1 25

The Last War Trail 1 25
LOG CABIN SERIES. By Edward S. Ellis. 3
vols., 12mo. Fully illustrated. Cloth, extra,
printed in colors. In box $3 75

Lost Trail 1 25
Camp-Fire and Wigwam 1 25
Footprints in the Forest 1 25

WYOMING SERIES. By Edward S. Ellis. 3

vols., 12mo. Fully illustrated. Cloth, extra,
printed in colors. In box $3 75

Wyoming 1 25
Storm Mountain 1 25
Cabin in the Clearing 1 25

New Books by Edward S. Ellis.

Through Forest and Fire. 12mo. Cloth $1 25
On the Trail of the Moose. 12mo. Cloth 1 25
 By C. A. Stephens.
Rare books for boys—bright, breezy, wholesome and instructive;
full of adventure and incident, and information upon natural history.
They blend instruction with amusement—contain much useful and
valuable information upon the habits of animals, and plenty of
adventure, fun and jollity.

CAMPING OUT SERIES. By C. A.

Stephens. 6 vols., 12mo. Fully illustrated.
Cloth, extra, printed in colors. In box $7 50
Camping Out. As recorded by "Kit" 1 25

Left on Labrador; or The Cruise of the

Schooner Yacht "Curfew." As recorded by
"Wash" 1 25

Off to the Geysers; or, The Young Yachters

in Iceland. As recorded by "Wade" 1 25

Lynx Hunting. From Notes by the author of

"Camping Out" 1 25
Fox Hunting. As recorded by "Raed" 1 25
On the Amazon; or, The Cruise of the
"Rambler." As recorded by "Wash" 1 25

By J. T. Trowbridge.
 These stories will rank among the best of Mr. Trowbridge's books
for the young—and he has written some of the best of our juvenile
literature.

JACK HAZARD SERIES. By J. T.

Trowbridge. 6 vols., 12mo. Fully
Illustrated. Cloth, extra, printed in colors.
In box $7 50
 *** END OF THE PROJECT GUTENBERG EBOOK THE YOUNG
CIRCUS RIDER; OR, THE MYSTERY OF ROBERT RUDD ***

Updated editions will replace the previous one—the old editions will
be renamed.

Creating the works from print editions not protected by U.S.

copyright law means that no one owns a United States copyright in
these works, so the Foundation (and you!) can copy and distribute it
in the United States without permission and without paying
copyright royalties. Special rules, set forth in the General Terms of
Use part of this license, apply to copying and distributing Project
Gutenberg™ electronic works to protect the PROJECT GUTENBERG™
concept and trademark. Project Gutenberg is a registered trademark,
and may not be used if you charge for an eBook, except by following
the terms of the trademark license, including paying royalties for use
of the Project Gutenberg trademark. If you do not charge anything
for copies of this eBook, complying with the trademark license is
very easy. You may use this eBook for nearly any purpose such as
creation of derivative works, reports, performances and research.
Project Gutenberg eBooks may be modified and printed and given
away—you may do practically ANYTHING in the United States with
eBooks not protected by U.S. copyright law. Redistribution is subject
to the trademark license, especially commercial redistribution.

START: FULL LICENSE

THE FULL PROJECT GUTENBERG LICENSE
 PLEASE READ THIS BEFORE YOU DISTRIBUTE OR USE THIS WORK

To protect the Project Gutenberg™ mission of promoting the free

distribution of electronic works, by using or distributing this work (or
any other work associated in any way with the phrase “Project
Gutenberg”), you agree to comply with all the terms of the Full
Project Gutenberg™ License available with this file or online at
www.gutenberg.org/license.

Section 1. General Terms of Use and

Redistributing Project Gutenberg™
electronic works
1.A. By reading or using any part of this Project Gutenberg™
electronic work, you indicate that you have read, understand, agree
to and accept all the terms of this license and intellectual property
(trademark/copyright) agreement. If you do not agree to abide by all
the terms of this agreement, you must cease using and return or
destroy all copies of Project Gutenberg™ electronic works in your
possession. If you paid a fee for obtaining a copy of or access to a
Project Gutenberg™ electronic work and you do not agree to be
bound by the terms of this agreement, you may obtain a refund
from the person or entity to whom you paid the fee as set forth in
paragraph 1.E.8.

1.B. “Project Gutenberg” is a registered trademark. It may only be

used on or associated in any way with an electronic work by people
who agree to be bound by the terms of this agreement. There are a
few things that you can do with most Project Gutenberg™ electronic
works even without complying with the full terms of this agreement.
See paragraph 1.C below. There are a lot of things you can do with
Project Gutenberg™ electronic works if you follow the terms of this
agreement and help preserve free future access to Project
Gutenberg™ electronic works. See paragraph 1.E below.
1.C. The Project Gutenberg Literary Archive Foundation (“the
Foundation” or PGLAF), owns a compilation copyright in the
collection of Project Gutenberg™ electronic works. Nearly all the
individual works in the collection are in the public domain in the
United States. If an individual work is unprotected by copyright law
in the United States and you are located in the United States, we do
not claim a right to prevent you from copying, distributing,
performing, displaying or creating derivative works based on the
work as long as all references to Project Gutenberg are removed. Of
course, we hope that you will support the Project Gutenberg™
mission of promoting free access to electronic works by freely
sharing Project Gutenberg™ works in compliance with the terms of
this agreement for keeping the Project Gutenberg™ name associated
with the work. You can easily comply with the terms of this
agreement by keeping this work in the same format with its attached
full Project Gutenberg™ License when you share it without charge
with others.

1.D. The copyright laws of the place where you are located also
govern what you can do with this work. Copyright laws in most
countries are in a constant state of change. If you are outside the
United States, check the laws of your country in addition to the
terms of this agreement before downloading, copying, displaying,
performing, distributing or creating derivative works based on this
work or any other Project Gutenberg™ work. The Foundation makes
no representations concerning the copyright status of any work in
any country other than the United States.

1.E. Unless you have removed all references to Project Gutenberg:

1.E.1. The following sentence, with active links to, or other

immediate access to, the full Project Gutenberg™ License must
appear prominently whenever any copy of a Project Gutenberg™
work (any work on which the phrase “Project Gutenberg” appears,
or with which the phrase “Project Gutenberg” is associated) is
accessed, displayed, performed, viewed, copied or distributed:
 This eBook is for the use of anyone anywhere in the United
States and most other parts of the world at no cost and with
almost no restrictions whatsoever. You may copy it, give it
away or re-use it under the terms of the Project Gutenberg
License included with this eBook or online at
www.gutenberg.org. If you are not located in the United
States, you will have to check the laws of the country where
you are located before using this eBook.

1.E.2. If an individual Project Gutenberg™ electronic work is derived

from texts not protected by U.S. copyright law (does not contain a
notice indicating that it is posted with permission of the copyright
holder), the work can be copied and distributed to anyone in the
United States without paying any fees or charges. If you are
redistributing or providing access to a work with the phrase “Project
Gutenberg” associated with or appearing on the work, you must
comply either with the requirements of paragraphs 1.E.1 through
1.E.7 or obtain permission for the use of the work and the Project
Gutenberg™ trademark as set forth in paragraphs 1.E.8 or 1.E.9.

1.E.3. If an individual Project Gutenberg™ electronic work is posted

with the permission of the copyright holder, your use and distribution
must comply with both paragraphs 1.E.1 through 1.E.7 and any
additional terms imposed by the copyright holder. Additional terms
will be linked to the Project Gutenberg™ License for all works posted
with the permission of the copyright holder found at the beginning
of this work.

1.E.4. Do not unlink or detach or remove the full Project

Gutenberg™ License terms from this work, or any files containing a
part of this work or any other work associated with Project
Gutenberg™.

1.E.5. Do not copy, display, perform, distribute or redistribute this

electronic work, or any part of this electronic work, without
prominently displaying the sentence set forth in paragraph 1.E.1
with active links or immediate access to the full terms of the Project
Gutenberg™ License.

1.E.6. You may convert to and distribute this work in any binary,
compressed, marked up, nonproprietary or proprietary form,
including any word processing or hypertext form. However, if you
provide access to or distribute copies of a Project Gutenberg™ work
in a format other than “Plain Vanilla ASCII” or other format used in
the official version posted on the official Project Gutenberg™ website
(www.gutenberg.org), you must, at no additional cost, fee or
expense to the user, provide a copy, a means of exporting a copy, or
a means of obtaining a copy upon request, of the work in its original
“Plain Vanilla ASCII” or other form. Any alternate format must
include the full Project Gutenberg™ License as specified in
paragraph 1.E.1.

1.E.7. Do not charge a fee for access to, viewing, displaying,

performing, copying or distributing any Project Gutenberg™ works
unless you comply with paragraph 1.E.8 or 1.E.9.

1.E.8. You may charge a reasonable fee for copies of or providing

access to or distributing Project Gutenberg™ electronic works
provided that:

• You pay a royalty fee of 20% of the gross profits you derive
from the use of Project Gutenberg™ works calculated using the
method you already use to calculate your applicable taxes. The
fee is owed to the owner of the Project Gutenberg™ trademark,
but he has agreed to donate royalties under this paragraph to
the Project Gutenberg Literary Archive Foundation. Royalty
payments must be paid within 60 days following each date on
which you prepare (or are legally required to prepare) your
periodic tax returns. Royalty payments should be clearly marked
as such and sent to the Project Gutenberg Literary Archive
Foundation at the address specified in Section 4, “Information
 about donations to the Project Gutenberg Literary Archive
Foundation.”

• You provide a full refund of any money paid by a user who

notifies you in writing (or by e-mail) within 30 days of receipt
that s/he does not agree to the terms of the full Project
Gutenberg™ License. You must require such a user to return or
destroy all copies of the works possessed in a physical medium
and discontinue all use of and all access to other copies of
Project Gutenberg™ works.

• You provide, in accordance with paragraph 1.F.3, a full refund of

any money paid for a work or a replacement copy, if a defect in
the electronic work is discovered and reported to you within 90
days of receipt of the work.

• You comply with all other terms of this agreement for free
distribution of Project Gutenberg™ works.

1.E.9. If you wish to charge a fee or distribute a Project Gutenberg™

electronic work or group of works on different terms than are set
forth in this agreement, you must obtain permission in writing from
the Project Gutenberg Literary Archive Foundation, the manager of
the Project Gutenberg™ trademark. Contact the Foundation as set
forth in Section 3 below.

1.F.

1.F.1. Project Gutenberg volunteers and employees expend

considerable effort to identify, do copyright research on, transcribe
and proofread works not protected by U.S. copyright law in creating
the Project Gutenberg™ collection. Despite these efforts, Project
Gutenberg™ electronic works, and the medium on which they may
be stored, may contain “Defects,” such as, but not limited to,
incomplete, inaccurate or corrupt data, transcription errors, a
copyright or other intellectual property infringement, a defective or
damaged disk or other medium, a computer virus, or computer
codes that damage or cannot be read by your equipment.

1.F.2. LIMITED WARRANTY, DISCLAIMER OF DAMAGES - Except for

the “Right of Replacement or Refund” described in paragraph 1.F.3,
the Project Gutenberg Literary Archive Foundation, the owner of the
Project Gutenberg™ trademark, and any other party distributing a
Project Gutenberg™ electronic work under this agreement, disclaim
all liability to you for damages, costs and expenses, including legal
fees. YOU AGREE THAT YOU HAVE NO REMEDIES FOR
NEGLIGENCE, STRICT LIABILITY, BREACH OF WARRANTY OR
BREACH OF CONTRACT EXCEPT THOSE PROVIDED IN PARAGRAPH
1.F.3. YOU AGREE THAT THE FOUNDATION, THE TRADEMARK
OWNER, AND ANY DISTRIBUTOR UNDER THIS AGREEMENT WILL
NOT BE LIABLE TO YOU FOR ACTUAL, DIRECT, INDIRECT,
CONSEQUENTIAL, PUNITIVE OR INCIDENTAL DAMAGES EVEN IF
YOU GIVE NOTICE OF THE POSSIBILITY OF SUCH DAMAGE.

1.F.3. LIMITED RIGHT OF REPLACEMENT OR REFUND - If you

discover a defect in this electronic work within 90 days of receiving
it, you can receive a refund of the money (if any) you paid for it by
sending a written explanation to the person you received the work
from. If you received the work on a physical medium, you must
return the medium with your written explanation. The person or
entity that provided you with the defective work may elect to provide
a replacement copy in lieu of a refund. If you received the work
electronically, the person or entity providing it to you may choose to
give you a second opportunity to receive the work electronically in
lieu of a refund. If the second copy is also defective, you may
demand a refund in writing without further opportunities to fix the
problem.

1.F.4. Except for the limited right of replacement or refund set forth
in paragraph 1.F.3, this work is provided to you ‘AS-IS’, WITH NO
OTHER WARRANTIES OF ANY KIND, EXPRESS OR IMPLIED,
INCLUDING BUT NOT LIMITED TO WARRANTIES OF
MERCHANTABILITY OR FITNESS FOR ANY PURPOSE.

1.F.5. Some states do not allow disclaimers of certain implied

warranties or the exclusion or limitation of certain types of damages.
If any disclaimer or limitation set forth in this agreement violates the
law of the state applicable to this agreement, the agreement shall be
interpreted to make the maximum disclaimer or limitation permitted
by the applicable state law. The invalidity or unenforceability of any
provision of this agreement shall not void the remaining provisions.

1.F.6. INDEMNITY - You agree to indemnify and hold the Foundation,

the trademark owner, any agent or employee of the Foundation,
anyone providing copies of Project Gutenberg™ electronic works in
accordance with this agreement, and any volunteers associated with
the production, promotion and distribution of Project Gutenberg™
electronic works, harmless from all liability, costs and expenses,
including legal fees, that arise directly or indirectly from any of the
following which you do or cause to occur: (a) distribution of this or
any Project Gutenberg™ work, (b) alteration, modification, or
additions or deletions to any Project Gutenberg™ work, and (c) any
Defect you cause.

Section 2. Information about the Mission

of Project Gutenberg™
Project Gutenberg™ is synonymous with the free distribution of
electronic works in formats readable by the widest variety of
computers including obsolete, old, middle-aged and new computers.
It exists because of the efforts of hundreds of volunteers and
donations from people in all walks of life.

Volunteers and financial support to provide volunteers with the

assistance they need are critical to reaching Project Gutenberg™’s
goals and ensuring that the Project Gutenberg™ collection will
remain freely available for generations to come. In 2001, the Project
Gutenberg Literary Archive Foundation was created to provide a
secure and permanent future for Project Gutenberg™ and future
generations. To learn more about the Project Gutenberg Literary
Archive Foundation and how your efforts and donations can help,
see Sections 3 and 4 and the Foundation information page at
www.gutenberg.org.

Section 3. Information about the Project

Gutenberg Literary Archive Foundation
The Project Gutenberg Literary Archive Foundation is a non-profit
501(c)(3) educational corporation organized under the laws of the
state of Mississippi and granted tax exempt status by the Internal
Revenue Service. The Foundation’s EIN or federal tax identification
number is 64-6221541. Contributions to the Project Gutenberg
Literary Archive Foundation are tax deductible to the full extent
permitted by U.S. federal laws and your state’s laws.

The Foundation’s business office is located at 809 North 1500 West,

Salt Lake City, UT 84116, (801) 596-1887. Email contact links and up
to date contact information can be found at the Foundation’s website
and official page at www.gutenberg.org/contact

Section 4. Information about Donations to

the Project Gutenberg Literary Archive
Foundation
Project Gutenberg™ depends upon and cannot survive without
widespread public support and donations to carry out its mission of
increasing the number of public domain and licensed works that can
be freely distributed in machine-readable form accessible by the
widest array of equipment including outdated equipment. Many
small donations ($1 to $5,000) are particularly important to
maintaining tax exempt status with the IRS.

The Foundation is committed to complying with the laws regulating

charities and charitable donations in all 50 states of the United
States. Compliance requirements are not uniform and it takes a
considerable effort, much paperwork and many fees to meet and
keep up with these requirements. We do not solicit donations in
locations where we have not received written confirmation of
compliance. To SEND DONATIONS or determine the status of
compliance for any particular state visit www.gutenberg.org/donate.

While we cannot and do not solicit contributions from states where

we have not met the solicitation requirements, we know of no
prohibition against accepting unsolicited donations from donors in
such states who approach us with offers to donate.

International donations are gratefully accepted, but we cannot make

any statements concerning tax treatment of donations received from
outside the United States. U.S. laws alone swamp our small staff.

Please check the Project Gutenberg web pages for current donation
methods and addresses. Donations are accepted in a number of
other ways including checks, online payments and credit card
donations. To donate, please visit: www.gutenberg.org/donate.

Section 5. General Information About

Project Gutenberg™ electronic works
Professor Michael S. Hart was the originator of the Project
Gutenberg™ concept of a library of electronic works that could be
freely shared with anyone. For forty years, he produced and
distributed Project Gutenberg™ eBooks with only a loose network of
volunteer support.
Project Gutenberg™ eBooks are often created from several printed
editions, all of which are confirmed as not protected by copyright in
the U.S. unless a copyright notice is included. Thus, we do not
necessarily keep eBooks in compliance with any particular paper
edition.

Most people start at our website which has the main PG search
facility: www.gutenberg.org.

This website includes information about Project Gutenberg™,

including how to make donations to the Project Gutenberg Literary
Archive Foundation, how to help produce our new eBooks, and how
to subscribe to our email newsletter to hear about new eBooks.
Welcome to our website – the ideal destination for book lovers and
knowledge seekers. With a mission to inspire endlessly, we offer a
vast collection of books, ranging from classic literary works to
specialized publications, self-development books, and children's
literature. Each book is a new journey of discovery, expanding
knowledge and enriching the soul of the reade

Our website is not just a platform for buying books, but a bridge
connecting readers to the timeless values of culture and wisdom. With
an elegant, user-friendly interface and an intelligent search system,
we are committed to providing a quick and convenient shopping
experience. Additionally, our special promotions and home delivery
services ensure that you save time and fully enjoy the joy of reading.

Let us accompany you on the journey of exploring knowledge and

personal growth!

ebookluna.com