(eBook PDF) Essentials of Ecology 4th Edition by Michael Begon

(eBook PDF) Essentials of Ecology 4th Edition by Michael Begon
(eBook PDF) Essentials of Ecology 4th Edition by Michael Begon
(eBook PDF) Essentials of Ecology 4th Edition by Michael Begon

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3. vi
Contents
Preface xi
Acknowledgments xiii
Part 1 Introduction 1
Chapter 1 Ecology and how to do it 3
1.1 What is ecology? 4
1.2 Scales, diversity of approaches,
and rigor 7
Questions of scale 7
The diversity of ecological evidence 10
Statistics and scientific rigor 12
1.3 Ecology in practice 15
Successions on old fields in Minnesota:
a study in time and space 15
Hubbard Brook: a long-term commitment to
study at the ecosystem scale 18
Canada’s Experimental Lakes Area: decades
of exploring the consequences of human
activities on lakes 20
An introduction of an exotic fish species
to New Zealand: investigation on multiple
biotic scales 21
Why Asian vultures were heading
for extinction: The value of a
modeling study 24
Summary 26
Review questions 27
Chapter 2 Ecology’s evolutionary
backdrop 28
2.1 Evolution by natural selection 29
2.2 Evolution within species 32
Geographical variation within
species 32
Variation within a species with ­man-made
selection pressures 37
Evolution and coevolution 39
2.3 The ecology of speciation 39
What do we mean by a ‘species’? 39
Allopatric speciation 40
Sympatric speciation? 44
2.4 The effects of climatic change
on the evolution and distribution
of species 46
2.5 Continental drift, parallel and
convergent evolution 49
2.6 Conclusion 52
Summary 52
Review Questions 53
Part 2 Conditions and Resources 55
Chapter 3 Physical conditions and
the availability of
resources 57
3.1 Environmental conditions 58
What do we mean by ‘harsh,’ ‘benign,’
and ‘extreme’? 58
Effects of conditions 59
Conditions as stimuli 61
The effects of conditions on interactions
between organisms 63
Responses by sedentary organisms 64
Animal responses to environmental
temperature 64
Microorganisms in extreme
environments 68
3.2 Resources for photosynthetic
organisms 68
Solar radiation 69
Water 72
Nutrients 75
Carbon dioxide 75
3.3 Heterotrophs and their resources 80
Nutritional needs and provisions 80
Defense 83
3.4 Effects of intraspecific competition for resources
 86

4. Contents  ■  vii  
3.5 Conditions, resources, and the ecological
niche 88
Summary 89
Review questions 89
Chapter 4 Climate and the world’s
biomes 91
4.1 The world’s climate 92
Redistribution of heat through
atmospheric movement 93
Ocean currents and the redistribution
of heat 95
4.2 Terrestrial biomes 95
Biomes and convergent evolution 99
Tropical rain forest 99
Savanna 104
Temperate grasslands 105
Desert 105
Temperate forest 106
Boreal forest (taiga) 106
Tundra 107
The future distribution of terrestrial
biomes 108
4.3 Aquatic ecosystems on the
continents 108
Streams and rivers 108
Lakes and ponds 112
Wetlands 113
4.4 Ocean biomes 113
The deep ocean 114
Subtropical gyres 115
Coastal upwelling systems 116
Broad continental shelves 117
Nearshore coastal marine
ecosystems 117
Summary 119
Review questions 120
Part 3 Individuals and
Populations 121
Chapter 5 Birth, death, and
movement 123
5.1 Populations, individuals, births
and deaths 124
What is an individual? 126
Counting individuals, births, and deaths 126
5.2 Life cycles 128
Life cycles and reproduction 128
Annual life cycles 129
Longer life cycles 130
5.3 Monitoring birth and death: life tables and fecundity schedules
 133
Cohort life tables 134
Life tables for populations with
overlapping generations 137
A classification of survivorship curves 138
5.4 Dispersal and migration 138
Dispersal determining abundance 140
The role of migration 142
5.5 The impact of intraspecific competition on populations
 142
Patterns of population growth 143
5.6 Life history patterns 147
Summary 152
Review Questions 152
Chapter 6 Interspecific competition 154
6.1 Ecological effects of interspecific competition
 155
Competition amongst phytoplankton
for phosphorus 155
Coexistence and exclusion of competing
salmonid fishes 156
Some general observations 157
Coexistence of competing diatoms 158
Coexistence of competing birds 159
Competition between unrelated
species 159
The competitive exclusion principle 160
Environmental heterogeneity 165
6.2 Evolutionary effects of interspecific competition
 169
Character displacement and ecological
release in the Indian mongoose 169
Character displacement in Canadian
sticklebacks 170

5. viii  ■  Contents
Evolution in action: selection on
microorganisms 170
6.3 Interspecific competition and community
structure 172
Limiting resources and the regulation of diversity in phytoplankton
communities 172
Niche complementarity amongst anem-
one fish in Papua New Guinea 172
Species separated in space or in time 174
Spatial separation in trees and tree-root
fungi 175
Temporal separation in mantids and
tundra plants 175
6.4 How significant is interspecific competition in practice?
 176
The prevalence of current competition 177
Competition or mere chance? 180
Summary 183
Review questions 184
Chapter 7 Predation, grazing, and
disease 185
7.1 What do we mean by predation? 186
7.2 Prey fitness and abundance 187
7.3 The subtleties of predation 189
Interactions with other factors 190
Compensation and defense by
individual prey 191
From individual prey to prey
populations 192
7.4 Predator behavior: foraging and
transmission 195
Foraging behavior 197
7.5 Population dynamics of predation 199
Underlying dynamics of predator–prey
interactions: a tendency to cycle 199
Predator–prey cycles in
practice 203
Disease dynamics and cycles 204
Crowding 207
Predators and prey in patches 209
7.6 Predation and community structure 211
Summary 214
Review questions 215
Chapter 8 Molecular and evolutionary
ecology 216
8.1 Molecular ecology: ­differentiation within
and between species 217
Differentiation within species:
albatrosses 221
Differentiation between species: the red
wolf—species or hybrid? 223
8.2 Coevolutionary arms races 226
Coevolution 226
Insect–plant arms races 226
Coevolution of parasites and their
hosts 229
8.3 Mutualistic interactions 232
Mutualistic protectors 232
Farming crops or livestock 233
The dispersal of seeds and
pollen 235
Mutualistic gut inhabitants 236
Mycorrhizas 236
Fixation of atmospheric nitrogen in
mutualistic plants 237
Summary 240
Review questions 240
Part 4 Communities and
Ecosystems 243
Chapter 9 From populations to
­communities 245
9.1 Multiple determinants of the dynamics of
populations 246
Fluctuation or stability? 247
Determination and regulation of
abundance 248
Using k -value analysis 250
9.2 Dispersal, patches, and metapopulation dynamics
 255
9.3 Temporal patterns in community composition
 259
Founder-controlled and dominance-controlled
communities 259
Community succession 262
9.4 Food webs 269

6. Contents  ■  ix  
Indirect and direct effects 269
Population and community stability and
food web structure 274
Summary 280
Review questions 281
Chapter 10 Patterns in species
richness 282
10.1 Quantifying species richness and
diversity 283
10.2 Spatially varying factors influencing species richness
 285
Productivity and resource richness 286
Energy 288
Predation intensity 292
Spatial heterogeneity 294
Environmental harshness 295
10.3 Temporally varying factors influencing species richness
 296
Climatic variation and its absence 296
Disturbance 296
Environmental age: evolutionary time 297
10.4 Habitat area and ­remoteness: Island
biogeography 298
10.5 Gradients of species richness 303
Latitudinal gradients 303
Gradients with altitude and depth 304
Gradients during community
succession 306
Summary 307
Review questions 308
Chapter 11 The flux of energy and matter
through ecosystems 309
11.1 The role of energy in ecology 310
11.2 Geographic patterns in ­primary
productivity 311
11.3 Factors limiting ­terrestrial primary
productivity 312
11.4 Factors limiting aquatic primary productivity
 318
11.5 The fate of primary ­productivity:
grazing 325
11.6 The process of decomposition 330
11.7 The flux of matter through ecosystems 332
11.8 Nutrient budgets and cycling at the ecosystem scale
 334
Summary 338
Review questions 339
Part 5 Applied Issues in Ecology 341
Chapter 12 Global biogeochemical
cycles and their alteration by
humans 343
12.1 What is biogeochemistry? 344
12.2 The global carbon dioxide cycle 345
Understanding the carbon dioxide
sinks 349
Effects of ocean acidification 351
Will the terrestrial carbon dioxide sink
change in the future? 351
Can we reduce carbon dioxide
emissions? 354
12.3 The global methane cycle 354
The natural sources of methane 356
Anthropogenic sources of methane 357
Methane and the global climate
system 357
How do we reduce methane
emissions? 360
12.4 The nitrogen cycle at global
and regional scales 361
Human acceleration of the nitrogen
cycle 362
The ecological and human-health
costs of nitrogen 364
Regional variation in nitrogen pollution 365
How can we reduce nitrogen
pollution? 367
Summary 368
Review questions 368
Chapter 13 Conservation ecology 370
13.1 The need for conservation 371
13.2 Small populations 375
The classification of risk 375
Demographic risks associated with
small populations 375

7. x  ■  Contents
Genetic problems in small populations 376
Habitat reduction 378
13.3 Threats to biodiversity 378
Overexploitation 378
Habitat disruption 379
Global environmental change 382
Introduced and invasive species 383
Infectious disease 386
Combinations of risks and extinction
vortices 386
Chains of extinctions? 386
13.4 Conservation in practice 389
Population viability analysis 390
Protected areas 393
Selecting conservation areas 397
Collections of areas 400
13.5 Ecosystem services 401
Summary 404
Review questions 404
Chapter 14 The ecology of human
­population growth, disease,
and food supply 406
14.1 Human use of ecological resources 407
14.2 The human population problem 408
Population growth up to the present 408
Predicting the future 410
Two future inevitabilities 411
A global carrying capacity? 414
14.3 Ecology and human health 414
Loss of the ozone layer 414
Extreme events 415
Changing global patterns of infection 416
Emerging infectious diseases 418
14.4 Synthetic fertilizer and the intensification
of agriculture 418
14.5 Monocultures, pests, and pesticides in agriculture
 421
Chemical approaches to pest
control 422
Biological control 426
14.6 Global land use and other constraints
on ­continued intensification of
agriculture 428
14.7 Food from fisheries and aquaculture 434
Summary 439
Review questions 440
References R-1
Index I-1
Glossary G-1
(available online only: www.wiley.com/ college/begon)

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

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

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

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

14. Ecology and how to do it
Chapter
1
CHAPTER CONTENTS
1.1
What is ecology?
1.2 Scales, diversity of approaches, and rigor
1.3 Ecology in practice
KEY CONCEPTS
After reading this chapter you will be able to:
• explain how ecologists seek to describe and
understand, and on the basis of their under-
standing, to predict, manage, mitigate, and
control
• describe the variety of spatial and temporal
scales on which ecological phenomena occur
• describe how ecologists use observations, field
and laboratory experiments, and mathematical
models to collect scientific evidence
© ElementalImaging/iSt
ockpho to

15. 4  ■   Part 1 Introduction
1.1 WHAT IS ECOLOGY?
We could answer the question ‘What is ecology?’ by
examining various definitions that
have been proposed and choosing
one as the best (Box 1.1). But while
definitions have conciseness and precision, and they
the earliest
ecologists
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.
are good at preparing you for an examination, they are
not so good at capturing the flavor and excitement of
ecology. There is a lot to be gained by replacing that
single question about a definition with a series of more
provocative ones: ‘What do ecologists do?’ ‘What are
ecologists interested in?’ and ‘Where did ecology
emerge from in the first place?’
1.1
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
Historical Landmarks

16. Ecology and how to do it Chapter 1  ■  5  
Ecology can lay claim to being the oldest science,
as the most primitive humans must have been ecolo-
gists of sorts, driven by the need to understand where
and when their food and their (nonhuman) enemies
were to be found. The earliest agriculturalists needed to
be even more sophisticated, with knowledge of how
to manage their domesticated sources of food. These
early ecologists, then, were applied ecologists, seeking
to understand the distribution, abundance, and produc-
tivity of organisms in order to apply that knowledge for
their own benefit. Applied ecologists today still have
many of the same interests: how to optimize the rate at
which food is collected from natural environments in a
sustainable way; how domesticated plants and animals
can best be managed so as to maximize rates of return;
how food organisms can be protected from their own
natural enemies; and how to control the populations of
pathogens and parasites that live on us.
In the last century or so, how-
ever, since ecologists have been
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),
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:
The bane of the recent development popularly
known as ecology has been a widespread feeling
that anyone can do ecological work, regardless of
a pure and applied
science
preparation. There is nothing . . . more erroneous
than this feeling.
On the other hand, the need for applied ecology
to be based on its pure counterpart was clear in the
introduction to Charles Elton’s (1927) Animal Ecology
(Figure 1.1):
Ecology is destined for a great future .  .  . The
tropical entomologist or mycologist or weed- controller will only be fulfilling his functions properly if he is first and foremost an ecologist.
In the intervening years, the coexistence of these
pure and applied threads has been maintained and built upon. Many applied sciences such as forestry, agronomy, and fisheries biology have contributed to the development of ecology and have seen their own
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.
courtesy Robert Elton
FIGURE 1.1 One of the great founders of ecology: Charles
Elton (1900–1991). Animal Ecology (1927) was his first book but
The Ecology of Invasions by Animals and Plants (1958) was equally influ-
ential. (After Breznak, 1975.)

17. 6  ■   Part 1 Introduction
development enhanced by ecological ideas and app­
roaches. All aspects of food and fiber gathering, pro-
duction, and protection have been involved. The
biological control of pests (the use of pests’ natural
enemies to control them) has a history going back at
least to the ancient Chinese but has seen a resurgence
of ecological interest since the shortcomings of chemi-
cal pesticides began to be widely apparent in the 1950s.
The ecology of pollution has been a growing concern
from around the same time and expanded further in
the 1980s and 1990s from local to regional and global
issues. The last few decades have also seen expansions
in both public interest and ecological input into the
conservation of endangered species and the biodiver-
sity of whole areas, the control of disease in humans as
well as many other species, and the potential conse-
quences of profound human-caused changes to the
global environment.
And yet, at the same time,
many fundamental problems of
ecology remain unanswered. To
what extent does competition for food determine
which species can coexist in a habitat? What role does
disease play in the dynamics of populations? Why are
there more species in the tropics than at the poles?
What is the relationship between soil productivity and
plant community structure? Why are some species
more vulnerable to extinction than others? Are wet-
lands net sources or sinks of greenhouse gas emission
to the atmosphere? And so on. Of course, unanswered
questions—if they are focused questions—are a symp-
tom of the health, not the weakness, of any science. But
ecology is not an easy science, and it has particular
subtlety and complexity, in part because ecology is
peculiarly confronted by ‘uniqueness’: millions of dif-
ferent species, countless billions of genetically distinct
individuals, all living and interacting in a varied and
ever-changing world. The beauty of ecology is that it
challenges us to develop an understanding of very
basic and apparent problems—in a way that recog-
nizes the uniqueness and complexity of all aspects of
nature – but seeks patterns and predictions within this
complexity rather than being swamped by it.
Let’s come back to the ques-
tion of what ecologists do. First
and foremost ecology is a science, and
ecologists therefore try to explain
and understand. Explanation can
be either ‘proximate’ or ‘ultimate,’ and ecologists are
interested in both. For example, the present distribu-
tion and abundance of a particular species of bird may
be ‘explained’ in terms of the physical environment
that the bird tolerates, the food that it eats, and the
unanswered
questions
understanding,
description,
­prediction, and
control
parasites and predators that attack it. This is a proxi-
mate explanation – an explanation in terms of what is going on ‘here and now.’ We can also ask how this bird came to have these properties that now govern its life. This question has to be answered by an expla-
nation in evolutionary terms; the ultimate explanation
of the present distribution and abundance of this bird lies in the ecological experiences of its ancestors (see Chapter 2).
In order to understand something, of course,
we must first have a description of whatever it is we wish to understand. Ecologists must therefore describe
before they explain. On the other hand, the most valu-
able descriptions are those carried out with a particular problem or ‘need for understanding’ in mind. Undirected description, carried out merely for its own sake, is often later found to have selected the wrong things and has little place in ecology—or any other science.
Ecologists also often try to predict. For exam-
ple, how will global warming affect the sequestra-
tion (storage) of carbon in natural ecosystems? Will warming reduce this storage, and therefore result in even more global warming since less carbon diox-
ide will be removed from the atmosphere? Often, ecologists are interested in what will happen to a population of organisms under a particular set of cir-
cumstances, and on the basis of these predictions to control, exploit or conserve the population. We try
to minimize the effects of locust plagues by predict-
ing when they are likely to occur and taking appro- priate action. We try to exploit crops most effectively by predicting when conditions will be favorable to the crop and unfavorable to its enemies. We try to preserve rare species by predicting the conservation policy that will enable us to do so. Some prediction and control can be carried out without deep explanation or understanding: it is not difficult to predict that the destruction of a woodland will eliminate woodland birds. But what if the woodland is not destroyed, but rather fragmented into distinct parts with suburbs or agricultural fields between them? What effect may this have on the woodland birds? Insightful predic- tions, precise predictions, and predictions of what will happen in unusual circumstances can be made only when we can also explain and understand what is going on.
This book is therefore about:
1
How ecological understanding is achieved.
2 What we do understand, and what we do not.
3 How ecological understanding can help us predict,
manage, mitigate, and control.

18. Ecology and how to do it Chapter 1  ■  7  
1.2 SCALES, DIVERSITY OF
APPROACHES, AND RIGOR
Ecology is a diverse discipline, and ecologists use a vast
array or tools and approaches. Later in this chapter, we
briefly give some examples of this diversity, but first
we elaborate on three general points:
• ecological phenomena occur at a variety of scales;
• ecological evidence comes from a variety of differ-
ent sources;
• ecology relies on truly scientific evidence.
Questions of scale
Ecology operates at a range of scales: time scales, spa-
tial scales, and ‘biological’ scales. It is important to
appreciate the breadth of these and how they relate
to one another.
Life is studied at a variety of
hierarchical levels, with much of biol-
ogy focused on levels from molecules,
to organelles, cells, tissues, organs, and whole organisms.
Ecologists study levels from individual organisms, to
populations, communities, ecosystems, and the global
biosphere (Figure 1.2).
• Populations are functioning groups of individual
organisms of the same species in a defined location.
• Communities consist of all the species populations
present in a defined location.
• Ecosystems include both the community of organisms
and the physical environment in which they exist.
• The biosphere is the totality of all of life interacting
with the physical environment at the scale of the
entire planet.
At the level of the organism, ecology deals primarily
with how individuals are affected by their environment
and with their physiological and behavioral responses
to the environment.
Population ecology stresses the
trends and fluctuations in the number of individual of a particular species at a particular time and place, as determined by the interactions of birth and death rates and the interactions between the populations them-
selves (such as predators and prey).
Community ecology
focuses on questions such as what controls the diversity of species of in a given area.
Ecosystem ecology strives to
understand the functioning of entire lakes, forests, wet-
lands, or other portions of the Earth in terms of energy and material inputs and outputs. Across all scales of biological hierarchy—including these ecological ones—
three generalities emerge.
the ‘biological’
scale
1 The properties observed at a particular level arise
out of the functioning of parts at the level below. For example, how a tissue functions is the result of the functioning of the cells in that tissue, and how an ecosystem functions is the result of the function- ing of the communities within it interacting with the physical environment.
2
In order to understand the mechanistic reasons that
a particular property is observed at any level of bio- logical organization, a scientist needs to look at the next lowest level of organization. To understand dysfunction in an individual organism, we must look at the functioning of the organs in that organ-
ism; and to understand the controls on birth rate in a population, we must look at reproduction in individual organisms.
3
However, properties observed at a given level of
organization may be predicted without fully under-
standing the functioning at lower levels. This third generality may seem to contradict the other two, but it does not. Consider an analogy from the physical sciences. As early as 1662, Boyle knew that when the pressure of a gas is doubled, its volume is halved, if temperature remains constant. This behavior of
the gas as a whole is the result of the interactions
of the gas molecules, yet Boyle’s law provided valu-
able predictive power for centuries, long before the concept of the molecule was developed. Today, physical chemists can indeed explain gas behavior based on understanding of the behavior of individ-
ual molecules, but the explanation is complex, and not even taught to most undergraduate college stu-
dents. Similarly, ecologists can predict patterns in ecosystems without understanding all of the details of the dynamics of constituent populations, and can predict patterns in populations without understand-
ing all of the details of the responses of individual organisms.
Within the living world, there
is no arena too small nor one so large that it does not have an ecol- ogy. Even the popular press talk increasingly about the ‘global ecosystem’, and there is no question that several ecological problems can be examined only at this very large scale. These include the relationships between ocean currents and fisheries, or between climate patterns and the distribution of deserts and tropical rain forests, or between elevated carbon dioxide in the atmosphere (from burning fossil fuels) and global climate change.
At the opposite extreme, an individual cell may
be the stage on which two populations of pathogens
a range of spatial
scales

19. 8  ■   Part 1 Introduction
Biosphere
Ecosystem
Community
Cells
Organism
Organs and tissues
THE REALM OF ECOLOGY OTHER LEVELS OF BIOLOGY
Population
Organelles
Molecules
B
r
y
k
a
y
l
o
Y
u
r
iy
/
S
h
u
tte
rs
to
c
k
Lag
u
n
a

D
e
s
i
g
n
/
S
c
i
e
n
c
e
S
o
u
rc
e
Lag
u
n
a

D
e
s
i
g
n
/
S
c
i
e
n
c
e
S
o
u
rc
e
Biophoto A
s
s
o
c
i
a
t
e
s
/
S
c
i
e
n
c
e

S
o
u
rc
e
m
i
c
r
o
_
p
h
o
t
o
/
i
S
t
o
c
k
p
h
o
to
Laguna D
e
s
i
g
n

/

S
c
i
e
n
c
e

S
o
u
rc
e
3D
C
li
n
i
c
/
G
e
t
t
y

I
m
a
g
e
s
NASA
Rola
n
d

B
i
r
k
e
/
P
e
t
e
r

A
r
n
o
ld
/
G
e
t
t
y

I
m
a
g
e
s
FIGURE 1.2 Ecology is studied at many hierarchical levels.

20. Ecology and how to do it Chapter 1  ■  9  
compete with one another for the resources that the
cell provides. At a slightly larger spatial scale, a ter-
mite’s gut is the habitat for bacteria, protozoans, and
other species (Figure 1.3) – a community whose diver-
sity is comparable to that of a tropical rain forest in
terms of the richness of organisms living there, the
variety of interactions in which they take part, and
indeed the extent to which we remain ignorant about
the species identity of many of the participants.
Between these extremes, different ecologists, or the
same ecologist at different times, may study the inhab-
itants of pools that form in small tree-holes, the tempo-
rary watering holes of the savannas, or the great lakes
and oceans; others may examine the diversity of fleas
on different species of birds, the diversity of birds in
different sized patches of woodland, or the diversity of
woodlands at different altitudes.
To some extent related to this
range of spatial scales, and to the
levels in the biological hierarchy,
ecologists also work on a variety of time scales.
Ecological succession – the successive and continuous
colonization of a site by certain species populations,
accompanied by the local extinction of others – may be
studied over a period from the deposition of a lump of
sheep dung to its decomposition (a matter of weeks),
from the abandonment of a patch of tropical rain for-
est cleared for slash-and-burn agriculture (years to
decades), or from the development of a new forest on
land wiped clean to bedrock by the retreat of a glacier
in the arctic or high mountains (centuries). Migration
a range of time
scales
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
for ecological investigation varies
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-
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
might be a cause for conservation concern—but one or
more decades of study may be needed to be sure that the
decline is more than just an expression of the random
ups and downs of ‘normal’ population dynamics. One
of the longest, continuously run ecological studies is
at the Hubbard Brook Experiment Forest in the White
Mountains of New Hampshire. Among other mea-
sures, Gene Liken and other scientists there have moni-
tored the acidity of rain since the early 1960s. In the
1960s, the rain was quite acidic (low pH: high hydro-
gen ion concentrations), and this was in fact one of
the earliest discoveries anywhere of the phenomenon
of acid rain. The long-term trend, though, has been for
precipitation to become less acidic over subsequent
decades (Figure 1.4); but we can observe this only
the need for long-
term studies
Shigeharu Moriya
FIGURE 1.3 The diverse community of a termite’s gut. Termites
can break down lignin and cellulose from wood because of their mutu-
alistic relationships (see Chapter 8) with a diversity of microbes that live
in their guts.
FIGURE 1.4 Hydrogen ion concentration in precipitation at the
Hubbard Brook Experimental Forest over time. Note the long-term trend
of decreasing concentration, indicating that the pH has been rising, and
the rain has become less acidic over time. However, analysis of periods
of only a few years in duration can show sharp increases or decreases
in the hydrogen ion concentration, and are quite misleading with regard
to the long-term trend (After Likens 2004).
1960
H
+
concentration in precipitation (µeq/L)
20
30
40
50
60
70
80
90
100
1965 1970 1975 1980 198 52 005
2000
19951990
We are able to observe a
long-term drop in rain acidity
only through studies over
many years.

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