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WILEY
Essentials of Ecology
Essentials of Ecology

| TH EDITION

MICHAEL BEGON
Department of Evolution, Ecology and Behaviour,
The University of Liverpool, Liverpool, UK

ROBERT W. HOWARTH
Department of Ecology & Evolutionary Biology,
Cornell University, Ithaca, New York, USA

COLIN R. TOWNSEND
Department of Zoology, University of Otago,
Dunedin, New Zealand

WILEY
VicE PRESIDENT AND EXECUTIVE PUBLISHER: Kaye Pace
Acqulisitions Eprror: Bonnie Roth
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ISBN 978-0-470-90913-3

Printed in the United States of America

10987654321
 Short Contents

Contents vi Chapter 8 Molecular and evolutionary

Preface xi ecology 216

Acknowledgments _ xiii
Part 4
Communities and Ecosystems 243
Part 1
Chapter9 From populations to
Introduction 1
communities 245
Chapter 1 Ecology andhowtodoit 3 Chapter 10 Patterns in species
Chapter 2 — Ecology's evolutionary richness 282
backdrop 28 Chapter 11 The flux of energy and matter
through ecosystems 309

Part 2
Conditions and Resources 55
Part 5
Applied Issues in Ecology 341
Chapter 3 Physical conditions and the
Chapter 12. Global biogeochemical
availability of resources 57
cycles and their alteration
Chapter 4 = Climate and the world's
by humans 343
biomes 91
Chapter 13. Conservation ecology 370
Chapter 14 The ecology of human

Part 3 population growth, disease,

Individuals and Populations 121 and food supply 406

Chapter 5 _ Birth, death, and References R-1

movement 123
Index I-1
Chapter 6 _ Interspecific
competition 154
Glossary G-1
Chapter 7 Predation, grazing, (available online only: www.wiley.com/
and disease 185 college/begon)
Preface xi What do we mean by a'species’? 39
Allopatric speciation 40
Acknowledgments xiii
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 todo it 3 2.5 Continental drift, parallel and
Ll 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
Part 2 Conditions and Resources 55
1.3 Ecology in practice 15
Successions on old fields in Minnesota: Chapter3 Physical conditions and
a study in time and space 15 the availability of
Hubbard Brook: a long-term commitment to resources 57
study at the ecosystem scale 18
3.1 Environmental conditions 58
Canada's Experimental Lakes Area: decades
What do we mean by ‘harsh, ‘benign;
of exploring the consequences of human
and ‘extreme’? 58
activities on lakes 20
Effects of conditions 59
An introduction of an exotic fish species
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 a 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
fel What do we mean by predation? 186 Review questions 240
7.2 Prey fitness and abundance 187
7.3 The subtleties of predation 189
Part4 Communities and
Interactions with other factors 190
Ecosystems 243
Compensation and defense by
individual prey 191 Chapter9 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
4 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 Foundercontrolled 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 Mm 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 Part5 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. Whatis 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
 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

References R-1
population growth, disease,
and food supply 406 Index +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- e 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 e ‘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 e ‘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
e ‘ECOncerns’ boxes highlight some of the applied
growth, disease, and food supply, takes an ecological
approach to examining the issues of the population problems in ecology, particularly those where there
is a social or political dimension (as there often is). In
problem, of human health, and of the sustainability of
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
e 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
e 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
e 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 Ecology and how to do it 3
2 Ecology's evolutionary
backdrop 28

Introduction
© Elementallmaging/Stockphoto
 es
lp
Ecology and how fo 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:
e explain how ecologists seek to describe and e 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

e describe the variety of spatial and temporal

scales on which ecological phenomena occur
 4 Parti Introduction

EF 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 IS ECOLOGY? 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 | Mecurlear single question about a definition with a series of more
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?’

Historical Landmarks

Definitions of ecology

Ecology (originally in German, Oekologie) 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 doit Chapter 1 a 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-
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),
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:
FIGURE 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 Parti 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-
many fundamental problems of 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
tion of what ecologists do. First only when we can also explain and understand what
and foremost ecology is a science, and is going on.
ecologists therefore try to explain 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 doit Chapter 1 a 7.

1.2 SCALES, DIVERSITY OF 1 The properties observed at a particular level arise

APPROACHES, AND RIGOR 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.
e ecological phenomena occur at a variety of scales; 2 In order to understand the mechanistic reasons that
e ecological evidence comes from a variety of differ- a particular property is observed at any level of bio-
ent sources; logical organization, a scientist needs to look at the
next lowest level of organization. To understand
e 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 standing the functioning at lower levels. This third
hierarchical levels, with much of biol- 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
e 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,
e 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-
e 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
e 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
trends and fluctuations in the number of individual of is no arena too small nor one so
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
 | Part 1 Introduction

Biosphere

Ecosystem

Community

Population

Organism

Organs and tissues

Organelles

Molecules

FIGURE 1.2 Ecology is studied at many hierarchical levels

 Ecology and how to doit Chapter 1 a 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
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.
Shigeharu Moriya

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
FIGURE 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-
rary watering holes of the savannas, or the great lakes
and oceans; others may examine the diversity of fleas
=
2
°
H* concentration in precipitation (yeq/L)

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
range of spatial scales, and to the 9 v4
levels in the biological hierarchy, 50 -}——

ecologists also work on a variety of time scales. 40

Ecological succession — the successive and continuous 30 x
colonization of a site by certain species populations, 20 1 1 1 i i 1
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
FIGURE 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),
10 ~~ Parti Introduction

because we have that long-term record. Observations same sort of data from many dif-
over periods of even 5 to 10 years at a time would be ferent sites. Consider the question
very misleading. whether the amount of nitrogen
This does not mean that all ecological studies pollution deposited onto the landscape in rain, snow,
need to last for 20 years — nor that every time an eco- dust, and gases affects the biodiversity of grassland
logical study is extended the answer changes. But it communities (nitrogen is a major component of acid
does emphasize the great value to ecology of the small rain). The extent of this nitrogen pollution varies
number of long-term investigations that have been car- greatly over the landscape of Europe, so we can test
ried out or are ongoing. the hypothesis that more nitrogen pollution lowers
biodiversity by comparing diversity in different grass-
The diversity of ecological evidence lands receiving different inputs of nitrogen pollu-
tion from the atmosphere (Figure 1.5). The diversity
Ecological evidence comes from a variety of different of forbs (broad-leaved herbs) is indeed lower when
sources and approaches. The principal tools are: nitrogen pollution is greater, but the diversity of
grasses increases with increasing nitrogen pollution.
e Observations, often of changes in abundance or sys-
tem functioning over either time or space, and often The scatter in the relationships is great, but the rela-
tionships are nonetheless significant. (We will discuss
involving comparisons across and between different
what we mean by “significant” later in this chapter.)
areas or systems.
The scatter is undoubtedly the result of other factors
e Experiments, including both those in the lab and in across the landscape - in addition to the nitrogen
the field. pollution — that might also affect diversity, such as
e Mathematical models that capture some component types of soil, differences in precipitation and other
of ecological interactions, function, and structure. climate variables, and other types of pollution and
disturbance.
Ecologists often combine two or more of these Rather than relying on this comparative obser-
approaches. For instance, they may use models or vational approach, we could conduct a manipulative
inferences from comparative observations across sys- field experiment to test the hypothesis that nitro-
tems to inform experiments, or they may use experi- gen pollution affects biodiversity. The result of one
ments and observations to calibrate models. such experiment shows a very tight and pronounced
Many ecological studies include effect of increasing nitrogen supply on biodiversity
careful observation and monitor- (expressed as species richness, the number of species)
ing in the natural environment, for after just 4 years of nitrogen addition (Figures 1.6).
instance, of the changing abundance of one or more This experiment has an advantage over the compara-
species over time, or over space, or both. In this way, tive observational study, in that other confounding
ecologists may establish patterns, for example, that red variables - soil type, climate, disturbance history -
grouse (birds shot for ‘sport’) exhibit regular cycles in are held constant between treatments and hence elim-
abundance peaking every 4 or 5 years. Documenting inated, and this probably explains the tighter rela-
the pattern does not provide explanation for the tionship between nitrogen and diversity. On the other
cause of the cycle, but it is a start toward under- hand, the experiment is conducted on just one type of
standing. A next step might be to development one or soil, with one type of climate and disturbance history,
more hypotheses to explain the pattern: for instance, and over a fairly limited period of time (4 years in this
perhaps the 4- to 5-year cycle is caused by a gradual case), and so the result may not fully apply to other
accumulation of parasitic worms in the grouse popu- areas and to longer time scales. Both manipulative
lations over this period of time. A manipulative field experiments and observations are critical to ecology,
experiment is One approach to test such a hypoth- and ecologists gain confidence in their understanding
esis, in this case by ridding the grouse of the parasites of nature when the two approaches lead to similar
and monitoring whether the 4- to 5-year population conclusions.
cycle persists. Treating the grouse for their parasites Why might nitrogen affect plant biodiversity? One
strongly dampens the population cycle, giving strong hypothesis is that the effect is one of fertilization, with
support to this hypothesis (Hudson, Dobson, & plants growing more as the nitrogen supply increases,
Newborn, 1998). and this leading to less diversity as species that grow
We can also use comparative field observations particularly well come to dominate the community and
to test hypotheses. That is, we explicitly compare the shade out other plants. A creative experiment gave some
 Ecologyandhowtodoit Chapter] mm II

fee ced brated

y less
is |

=
Grass richness as a proportion

Forb richness as a proportion

of species richness

of species richness

FIGURE 1.5 The relationship between deposi-

tion of nitrogen pollution from the atmosphere and
diversity of grasses (panel “a”) and forbs (panel “b”)
across a wide number of grasslands on acid soils in
Europe. The dots on the map show the sifes sampled.
The photo shows a typical grassland from the Ukraine
(After Stevens et al.,2011).

Perhaps less obviously, ecolo-

gists also often turn to laboratory
systems or to mathematical models
a designed to capture ecological pro-
3<
=
cesses. These have played a crucial role in the develop-
2 ment of ecology, and they are certain to continue to
o
2
° do so. Field experiments are almost inevitably costly
o

a
Qa
and difficult to carry out. Moreover, even if time and
expense were not issues, natural field systems may
simply be too complex to allow us to tease apart the
0 50 100 150 200 250 300 consequences of the many different processes that
Nitrogen addition (kg N ha™ yr“) may be going on. Are the intestinal worms actually
capable of having an effect on reproduction or mor-
FIGURE 1.6 Experimental addition of nitrogen to a grassland in tality of individual grouse? How do light and nitrogen
Minnesota reduces species diversity. Reprinted from Huston (1997),
interact to regulate the growth rate of the various spe-
based on data in Tilman (1996).
cies in a grassland ecosystem? Controlled laboratory
experiments are often the best way to provide answers
preliminary support for this hypothesis: when artificial to specific questions that are key parts of an overall
light was supplied to the shaded plants of the understory explanation of the complex situation in the field
in nitrogen fertilized plots, biodiversity was maintained Of course, the complexity of natural ecological
despite the higher nitrogen (Hautier et al., 2009). communities may simply make it inappropriate for an
12) 3 @ Parti Introduction

ecologist to dive straight into them level of confidence to our conclusions. Ecology, like all
in search of understanding. We may science, is a search not for statements that have been
wish to explain the structure and ‘proved to be true’ but for conclusions in which we can
dynamics of a particular community of 20 animal and be confident.
plant species comprising various competitors, preda- What distinguishes science -
tors, parasites, and so on (relatively speaking, a com- what makes science rigorous — is
munity of remarkable simplicity). But we have little that it is based not simply on asser-
hope of doing so unless we already have some basic tions, but rather on conclusions
understanding of even simpler communities of just one resulting from investigations that
predator and one prey species, or two competitors, or test specific hypotheses, and to which we can attach a
(especially ambitious) two competitors that also share level of confidence, measured on an agreed-upon scale.
a common predator. For this, it is usually most appro- Statistical analyses are car-
priate to construct, for our own convenience, simple ried out after data have been col-
laboratory systems that can act as benchmarks or lected, and they help us to interpret
jumping-off points in our search for understanding. those data. Really good science, though, requires
What is more, you have only forethought. Ecologists, like all scientists, must know
to ask anyone who has tried to rear what they are doing, and why they are doing it, while
caterpillar eggs, or take a cohort of they are doing it. Ecologists must plan, so as to be
shrub cuttings through to maturity, to discover that even confident that they will collect the right kind of data,
the simplest ecological communities may not be easy to and a sufficient amount of data, to address the ques-
maintain or keep free of unwanted pathogens, predators, tion they hope to answer. As discussed in Box 1.2,
or competitors. Nor is it necessarily possible to construct more data are required to obtain statistically signifi-
precisely the particular, simple, artificial community that cant results when the relationship being tested is a
interests you; nor to subject it to precisely the conditions weak one, or when the relationship is confounded by
or the perturbation of interest. In many cases, therefore, other factors, as is likely the case for the relationship
there is much to be gained from the analysis of math- between nitrogen deposition and diversity illustrated
ematical models of ecological communities: constructed in Figure 1.5.
and manipulated according to the ecologist’s design. Many ecological field experi-
On the other hand, although a major aim of sci- ments rely on a large number of
ence is to simplify, and thereby make it easier to under- replicates for each treatment, and
stand the complexity of the real world, ultimately it this increases the likelihood of obtaining statistically
is the real world that we are interested in. The worth significant results. For example, ecologists experimen-
of models and simple laboratory experiments must tally testing the effect of nitrogen deposition on plant
always be judged in terms of the light they throw on diversity in grasslands might have 8 different levels of
the working of more natural systems. They are a means nitrogen inputs, with 10 different plots for each treat-
to an end—never an end in themselves. Like all scien- ment (a total of 80 plots). However, replication can be
tists, ecologists need to ‘seek simplicity, but distrust it’ expensive and time consuming, particularly if the ecol-
(Whitehead, 1953). ogists include in the responses they monitor processes
that are difficult to measure. Determining the biomass
of the plants at the end of the experiment is relatively
Statistics and scientific rigor
simple (cutting, drying, and weighing); characterizing
For a scientist to take offense at some popular phrase the diversity of the community is more difficult, par-
or saying is to invite accusations of a lack of a sense of ticularly if the diversity is high with many species
humor. But it is difficult to remain calm in the face of potentially present; measuring the rate at which each
phrases such as ‘There are lies, damn lies and statistics’ species is assimilating nitrogen is far, far more difficult
or ‘You can prove anything with statistics.’ Statistics and time consuming. Most experimentalists feel a con-
are regularly misused, but more often in the public stant tug between having a large number of replicates
media and by those who may seek to manipulate pop- and keeping their experiments doable.
ular opinion, and rarely if ever in the scientific litera- As noted by David Schindler
ture. You should not mistrust statistics. Rather, you (1998), experiments can often
should understand their strengths and limitations. An involve a trade-off between realism
essential point: you cannot prove anything with statis- and replication. Smaller scale
tics. Rather, statistical analysis allows us to attach a experiments — such as small plots of grassland, or
 Ecology and how to doit Chapter 1 13

re

Interpreting probabilities

Ecologists need to know, as do any scientists dealing with sets of data, what conclusions can be drawn
from those data. Imagine we are interested in determining whether high abundances of a pest insect in
summer are associated with high temperatures the previous spring, and imagine we have data on summer
insect abundances and mean spring temperatures for each of a number of years. How do we use statisti-
cal analysis to conclude, with a stated degree of confidence, either than there is or is not a relationship
between the spring temperature and summer insect numbers?

Null hypotheses and P-values

To carry out a statistical test we first need a null hypothesis, which simply means in this case that there is no
association; that is, no association between insect abundance and temperature. The statistical test (stated
simply) then generates a probability (a P-value) of getting a data set like ours if the null hypothesis is correct.
Suppose the data were like those in Figure 1.7a.The probability generated by a statistical test of asso-
ciation on these data is P= 0.5 (equivalently 50%). This means that, if the null hypothesis really was correct
(no association), then 50% of studies like ours should generate just such a data set, or one even further
from the null hypothesis. We therefore could have no confidence in any claim that there was an association.
Suppose, however, that the data were like those in Figure 1.7b, where the P-value is 0.001 (0.1%).
This would mean that such a data set (or one even further from the null hypothesis) could be expected
in only 0.1% of similar studies if there was really no association. In other words, either something very
improbable has occurred, or there was an association between insect abundance and spring temperature.
Thus, since we do not expect highly improbable events to occur, we can have a high degree of confidence
in the claim that there was an association between abundance and temperature.

Significance testing
Both 50% and 0.01%, though, make things easy for us. Where, between the two, do we draw the line? There
is no absolute answer to this, but scientists and statisticians have established a convention in significance
testing, which says that if P is less than 0.05 (5%), written P < 0.05 (e.g., Figure 1.7d), then results are
described as ‘statistically significant’ and confidence can be placed in the effect being examined; whereas
if P> 0.05, then there is no statistical foundation for claiming the effect exists (e.g., Figure 1.7c).A further
elaboration of the convention offen describes results with P < 0.01 as ‘highly significant’

‘Insignificant’ results?
Some effects are naturally strong (there is a powerful association between people’s weight and their
height) and others are weak (the association between people's weight and their risk of heart disease is real
but weak, since weight is only one of many important factors). More data are needed to establish support
for a weak effect than for a strong one. Hence a P-value of greater than 0.05 (lack of statistical significance)
may mean one of two things in an ecological study:

1 There really is no effect of ecological importance.

2 The data are simply not good enough, or there are not enough of them, to support the effect even
though it exists, possibly because the effect itself is real but weak.

Throughout this book, then, studies of a wide range of types are described, and their results often
have P-values attached to them. Remember that statements like P < 0.05 and P < 0.01 mean that these
are studies where: (i) sufficient data have been collected to establish a conclusion in which we can be con-
fident; (ii) that confidence has been established by agreed means (statistical testing); and (iii) confidence
is being measured on an agreed and interpretable scale.
14 Part 1 Introduction

P=0.5, the scatter is great, P =0.001, allowing us to

and there is no reason to conclude that abundances
believe abundance is related are greater at higher
(a) to temperature. (b) temperatures.

Zs
—
.
o
Q
g 10 1 12 13 14 15
3
§£ P =0.1, hinting at a relationship P =0.04, which indicates a stronger
8 (c) between temperature and abundance. (d) relationship than in Fig. 1.7-c,
& 25 —_——t supporting the conclusion that
2 . abundances rise with temperature. a
s
2 cast
=z 20 ° Rs
° eT
-
i ° soriF «re
a ° gat" °
10 >=ae
in
° me
5 a6 = °
s es wer °
6 °°
oket 1 L eo | J
10 1 12: 13 14 15 10 VW 12 13 14 15
Mean spring temperature (°C)

FIGURE 1.7 The results from four hypothetical studies of the relationship between insect pest abundance in summer
and mean temperature the previous spring, In each case, the points are the data actually collected. Horizontal lines represent
the null hypothesis — that there is no association between abundance and temperature, and thus the best estimate of expected
insect abundance, irrespective of spring temperature, is the mean insect abundance overall. The second line is the line of best
fitto the data, which in each case offers some suggestion that abundance rises as temperature rises. However, whether we can
be confident in concluding that abundance does rise with temperature depends, as explained in the text, on statistical tests
applied to the data sets. (a) The suggestion of a relationship is weak (P = 0.5). There are no good grounds for concluding that
the true relationship differs from that supposed by the null hypothesis and no grounds for concluding that abundance is related
to temperature. (b) The relationship is strong (P = 0.001) and we can be confident in concluding that abundance increases
with temperature. (c) The results are suggestive (P= 0.1) but it would not be safe to conclude from them that abundance rises
with temperature, (d) The results are not vastly different from those in (c) but are powerful enough (P= 0.04, i.e.,P < 0.05) for
the conclusion that abundance rises with temperature to be considered safe.

Standard errors and confidence intervals

Another way in which our confidence in results is assessed is through reference to ‘standard errors; which
statistical tests offen allow to be attached either to mean values calculated from a set of observations or
to slopes of lines like those in Figure 1.7.These mean values and slopes can only ever be estimates of the
‘true’ mean value or slope, because they are calculated from data that are only a sample of all the imagin-
able items of data that could be collected. The standard error, then, sets a band around the estimated value
within which the true value can be expected to lie, with a given, stated probability. In particular, there is a
95% probability that the true mean lies within roughly two standard errors (2 SE) of the estimated mean;
we call this the 95% confidence interval.
Large standard errors (little confidence in the estimated value) can arise when data are, for whatever
reason, highly variable; but they may also be due to only a small data set having been collected. Standard
errors are smaller, and confidence in estimates greater, both when data are more consistent (less variable)
and when there are more data.
 Ecologyandhowtodoit Chapter] mm 15

50-gallon containers of pond water - make replication Lake trout in a reference

relatively easy, but this scale cannot capture all of the lake receiving no acid
complex interactions inherent in natural ecosystems. are healthy.

Schindler has devoted his career to whole-ecosystem

experiments, for instance, adding nutrients, acids, or
toxins to whole lakes and following the response of the
lake ecosystem. At this scale, replication is difficult at
best, and many whole-ecosystem experiments have not
used replicate treatments. An excellent example is the
Fisheries and Oceans Canada
work of Schindler and colleagues (1985), who added
sulfuric acid to one lake (no replication) over a period Lake trout
in the acidified
of many years, to determine what the consequences of lake are clearly underweight
in the years before they are
acid rain on lakes might be. To understand the effects finally gone.
of the acidification, they observed the lake for a
period of years before acidification, and then observed
the behavior of the acidified lake in comparison to
other nearby reference lakes as acidification was grad-
ually increased. They found remarkable effects at levels
of acidity far less than lab studies of the time had indi- Fisheries and Oceans Canada
cated were likely to be a problem. For instance, the
populations of crayfish and small minnows in the lake FIGURE 1.8 After several years of acidification, lake trout in the
experimentally acidified lake (bottom) are obviously less healthy than
declined, and this led to deleterious consequences
those from a nearby reference lake (top). This resulted not from a direct
for the top predator, the lake trout (Figure 1.8), and to
toxicity effect of the acid on the fish, but rather on less prey organisms
their eventual disappearance from the lake. Note also
available for the fish. Reprinted from Schindler et al. (1985).
that while the experimental treatment (acidification)
was not replicated, the scientists took a large number
of replicate samples for the response variables of inter- pond can tell us something about fish of that species in
est (for instance, numbers of crayfish) in both the acidi- ponds of that type, generally. In short, ecology relies on
fied and reference lakes over time, making statistically obtaining estimates from representative samples. This
significant conclusions possible. While at first glance it is elaborated in Box. 1.3.
may seem distasteful to purposely pollute a lake, this
whole-lake experiment clearly demonstrated more
severe effects of acid rain on lakes than had previously 1.3 ECOLOGY IN PRACTICE
been believed possible, and this led to pressure to
To discover the real problems faced by ecologists and
reduce acid rain, protecting a far greater number of
how they try to solve them, we consider some real
lakes than the one damaged in the experiment.
research programs in a little detail. Every chapter in
Ecologists often seek to draw
conclusions about overall groups this book will contain descriptions of similar studies,
of organisms or overall trends in but in the context of a systematic survey of the driv-
ing forces in ecology (Chapters 2-11) or of the appli-
processes or fluxes: what is the
cation of this knowledge to solve applied problems
trend in abundance over time of the crayfish in the
(Chapters 12-14). For now, we want to highlight some
lake experiment, described above? What is the birth
of the excitement and diversity of ecological studies,
rate of the bears in Yellowstone Park? What is the den-
giving you the flavor of our field. We have chosen the
sity of weeds in a wheat field? What is the rate of loss
following examples either because they are classic stud-
in streams of nitrogen from forests? In doing so, we can
only very rarely examine every individual in a group, ies that pioneered new approaches, or because they
exemplify the use of multiple approaches in addressing
or in the entire sampling area, or all the nitrogen in all
the water leaving a forest. We must therefore rely on fundamental ecological questions.
what we hope will be a representative sample from the
Successions on old fields in Minnesota:
group or habitat or flux. Indeed, even if we examined
a study in time and space
a whole group (we might examine every fish in a small
pond, say), we are likely to want to draw general con- Ecological succession is a concept familiar to you if
clusions from it: we might hope that the fish in ‘our’ you have simply taken a walk in open country — the
16 =~ Parti Introduction

es

Estimation: sampling, accuracy, and precision

In any sampling program, the aim is to;

1 obtain an estimate that is accurate and unbiased; that is, neither systematically too high nor too low as
a result of some flaw in the program.

2. obtain data that have as little variation as possible.

3 use the time, money, and human effort invested in the program as effectively as possible, because these
are always limited.

To understand the application of these goals, consider another hypothetical example: the density
of a particular weed (e.g., wild oat) in a wheat field. To prevent bias, each part of the field should have an
equal chance of being selected for sampling, so sampling units should be selected at random. We might,
for example, divide the field into a measured grid, pick points on the grid at random, and count the wild
oat plants within a 50 cm radius of the selected grid point. This unbiased method can be contrasted with
a plan to sample only weeds from between the rows of wheat plants, giving too high an estimate, or within
the rows, giving too low an estimate (Figure 1.9a).
What, though, if we suspect that the slope of a field affects weed density, and half the field slopes
fo the southeast and half to the southwest? The individual values from samples may then fall into two
groups a substantial distance apart on the density scale: high from the southwest slope; low (mostly zero)
from the southeast slope. The estimated mean weed density would be close to the true mean (it would
be accurate and unbiased), but the variation among samples would be large and the estimate therefore
imprecise (Figure 1.9b). If, however, we acknowledge the difference between the two slopes and treat them
separately from the outset, then we obtain means for each that have much smaller confidence intervals.

(a) (b) (°c)

Study Study Study Single study of SE and SW studied SE and SW studied
1 2 3 the whole field separately, then combined separately, then combined
——— | ae |
oe ee eccceces

SW slope
Weeds per m?—>

True mean True mean

Random Between Within Individual Estimate SW SE Combined _ Individual SW SE Combined

sample rowsonly rows only samples estimate estimate estimate samples estimate estimate estimate

FIGURE 1.9 The results of hypothetical programs to estimate weed density in a wheat field. (a) The three studies have equal
potential error or variation associated with their estimates (as shown by the “error bars”), but only the first (from a random sample)
is accurate. (b) In the first study, individual samples from different parts of the field (southeast and southwest) fall into two groups
(left); thus, the estimate, although accurate, is not precise (right).In the second study, separate estimates for southeast and southwest
are both accurate and precise - as is the estimate for the whole field obtained by combining them. (c) Following on from (b), most
sampling effort is directed to the southwest, reducing the confidence interval there, but with little effect on the confidence interval for
the southeast. The overall interval is therefore reduced: precision has been improved.
 Ecologyandhowtodoit Chapter] mm 17

idea that a newly created habitat, or one in which a successional sequence of plants that occur in fields in
disturbance has created an opening, will be inhabited, the years following abandonment, we could plan an
in turn, by a variety of species appearing and disap- artificial manipulation, under our control, in which a
pearing in some recognizably repeatable sequence. number of fields currently under cultivation were forc-
Widespread familiarity with the idea, however, does ibly abandoned and the communities in them sampled
not mean that we understand fully the processes that repeatedly into the future. (We would need a number
drive or fine-tune successions; yet developing such of fields because any single field might be atypical,
understanding is important not just because succession whereas several would allow us to calculate mean val-
is one of the fundamental forces structuring ecological ues for, say, the number of new species per year, and
communities, but also because human disturbance of place confidence intervals around those means.) But
natural communities has become ever more frequent the results of this experiment would take decades to
and profound. We need to know how communities accumulate. The natural experiment alternative, there-
may respond to, and hopefully recover from, such dis- fore, was to use the fact that records already exist of
turbance, and how we may aid that recovery. when many of the old fields were abandoned. This
One particular focus for the study of succession is what David Tilman and his team did. Figure 1.10
has been the old agricultural fields of the eastern and illustrates data from a group of 22 old fields surveyed
northern United States, abandoned as farmers moved in 1983, having been abandoned at various times
west in search of ‘fresh fields and pastures new.’ One between 1927 and 1982 (i.e., between 1 and 56 years
such site is now the Cedar Creek Natural History Area, previously). These can be treated as 22 ‘snapshots’ of
roughly 50 km north of Minneapolis, Minnesota. The the continuous process of succession in old fields at
area was first settled by Europeans in 1856 and was Cedar Creek in general, even though each field was
initially subject to logging. Clearing for cultivation itself only surveyed once.
then began about 1885, and land was first cultivated A number of the shifting balances during succes-
between 1900 and 1910. Now there are agricultural sion are clear from the figure as statistically significant
fields that are still under cultivation and others that trends. Over the 56 years, the cover of invader species
have been abandoned at various times since the mid- (mostly agricultural weeds) decreased (Figure 1.10a)
1920s. Cultivation led to depletion of nitrogen from while the cover of species from nearby prairies increased
soils that already were naturally poor in this important (Figure 1.10b): the natives reclaimed their land. Of
plant nutrient. more general applicability, the cover of annual species
Studies at Cedar Creek illus- decreased over time, while the cover of perennial spe-
trate the value of natural experi- a | cies increased (Figure 1.10c, d). Annual species - those
ments. A natural experiment is that complete a whole generation from seed to adult
not planned before hand, but rather ecologists take through to seeds again within a year — tend to be good
advantage of a situation where either natural events at increasing in abundance rapidly in relatively empty
(the retreat of a glacier, to evaluate how plants recol- habitats (the early stages of succession); whereas peren-
onize soils) or human-controlled events set the stage nials - those that live for several or many years and
to learn something about ecological processes. The may not reproduce in their early years — are slower to
Cedar Creek studies, begun only in the 1980s, take establish but more persistent once they do. Over time,
advantage of the abandonment of agricultural land nitrogen accumulates in the soil (Figure 1.10e). Such
beginning many decades earlier. To understand the data suggest that the change in nitrogen may drive some
18 Parti Introduction

s
Percent cover
Percent cover

0 10 20 30 40 50 60
Field age (years)
g

1000
Soil nitrogen (mg kg“)

800

600

400

0 10 20 30 40 50 60
Field age (years)

FIGURE 1.10 Twenty-two fields at different stages in an old-field succession were surveyed to generate the following trends with successional
stage (field age): (a) invader species decreased, (b) native prairie species increased, (c) annual species decreased, (d) perennial species increased,
and (e) soil nitrogen content increased. The best fit lines (see Box 1.2) are highly significant in every case (P < 0.01). (After Inouye et al., 1987.)

of the changes in plant community structure. Or con- Herb Bormann, has been doing precisely this in the
versely, perhaps the differences in nitrogen are the result Hubbard Brook Experimental Forest since the early
of the differences in the plant community. Hypotheses 1960s. In part because of the noted success of having
based on these observations have encouraged many long-term continuous data demonstrated at Hubbard
field manipulative experiments and sparked some lively Brook, the U.S. National Science Foundation has now
debates over the past few decades, topics we explore in for several decades supported a network of Long-Term
more detail later in this book. Ecological Research sites, in a variety of different types
of ecosystems. Currently, 26 such sites are funded
(http://www.lternet.edu/). The goal is to continue these
Hubbard Brook: a long-term commitment to
efforts indefinitely into the future, providing a critical
study at the ecosystem scale
laboratory for studying ecological processes and how
The Cedar Creek study took advantage of space-for- they change over time.
time substitutions, inferring information on ecological An early accomplishment of
processes that occur over decades (old field succession the Hubbard Brook team was to
in Minnesota) from current spatial patterns. For longer develop the small watershed tech-
time scales, this may be the only approach for under- nique to measure the input and out-
standing ecological processes and controls. At shorter put of chemicals from individual forest areas. Because
time scales, though, actual observations are possible. most of the chemical losses from forests such as those
As noted briefly above, a project started by Gene at Hubbard Brook (with shallow soils, underlain by
Likens together with many other colleagues, notably impermeable bedrock) are channeled through streams,
 Ecologyandhowtodoit Chapter] mm 19

TABLE 1.1 Annual chemical budgets for forested catchment areas at Hubbard Brook (kg ha-' yr-'). Inputs are for dissolved materials in
precipitation or in dryfall (gases or associated with particles falling from the atmosphere). Outputs are losses in stream water as dissolved material
plus particulate organic material in the streamflow. The source of the excess chemicals (where outputs exceeded inputs) was weathering of parent
rock and soil. The exception was nitrogen (as ammonium or nitrate ions) — less was exported than arrived in precipitation because of nitrogen
uptake in the forest (After Likens and Bormann 1994),

ry Ne 0) as (ert AY era No
Input 27 16.3 38.3 Tk 2.6 0.7 5

Output 0.4 87 48.6 Vee. 11.8 29 69

Net Change* +23 +76 — 10.3 -0.6 -9.2 -2.2 -54

*Net change is positive when the catchment gains matter and negative when it loses it.

a comparison of the chemistry of stream water with

that of incoming precipitation can reveal a lot about
the differential uptake and cycling of chemical ele-
ments by the terrestrial biota. These small watersheds
are often called catchments.
In their catchment studies, the Hubbard Brooks

US Forest Service, Northern Research Station

scientists measured (and continue to measure!) the
inputs of acids, nutrients, and other materials from
the atmosphere. They use a variety of tools, but the
simplest is to measure total precipitation (rain and
snow) and the concentrations of materials in the pre-
cipitation (measured in plastic buckets, which are only
open to the air when it is raining), as well as what falls
in other buckets at times of no rain (automatic sensors
move covers depending upon whether it is raining or
not). The scientists also measure what leaves the catch- FIGURE 1.11 The Hubbard Brook Experimental Forest. Note the
ments in streamflow, using carefully gauged streams experimental stream catchment from which all trees were removed, left
which provide a precise measure of water flow over of photo.
time and periodic sampling of the chemicals dissolved
in that stream water. Their early results (Table 1.1)
were very surprising for the time, and of great interest: loggers sometimes do in what is called a “clear cut”
for many substances, the export from the forest (Figure 1.11), and compared the response of export
exceeded inputs from the air (the result of weathering of materials in stream-water with that in another 5
of the soil and bedrock material). But for nitrogen reference watersheds. The overall export of dissolved
(both ammonium, NH,*, and nitrate, NO) the export inorganic substances from the disturbed catchment
from the ecosystem was less than the inputs: the forest rose to 13 times the normal rate (Figure 1.12). Two
was retaining a large amount of the nitrogen inputs. phenomena were responsible. First, the enormous
Likens and colleagues also
pioneered the idea of perform- large-s
al nfold
egesoa
ald
reduction in transpiring surfaces (leaves) on live
trees led to 40% more of the input precipitation
ing large-scale ecological experi- - experiment being exported again as water in the streams, carry-
ments at the catchment scale. ing with it more dissolved substances. Second, and
Hydrologists had done similar experiments before more importantly, as microbes decomposed organic
but had not explored the ecological ramifications. matter and made substances available in solution,
In the first of the ecologically oriented experiments, the trees no longer were there to assimilate these
the Hubbard Brook team completely cut down all materials; rather, they were flushed from the system
of the trees in one small catchment, as commercial with the exported water.
20 sg Part 1 Introduction

— Deforested catchment
—— Control catchment

2.0 a / LyANA
Concentration (mg I")

1.05

4.0 Kt
3.0
2.0 MY
1.0 |

80 NO.
60
40 -
20 FIGURE 1.12 Concentrations of ions in
4.0 - “¢- - stream water from the experimentally defor-
3.0 | ested watershed 2 and the control (unmanipu-

ot AW
lated) watershed 6 at Hubbard Brook. The timing
of deforestation is indicated by arrows. In each
case, there was a dramatic increase in export
JASONDJFMAMJJASONDJFMAMJJASONDJFMAM of the ions after deforestation. Note that the
eee IN nA oN YO NS v see
1965 1966 1967 1968 ‘nitrate’ axis has a break in it. (After Likens &
Date Bormann, 1975.)

The Hubbard Brook team has gone on to per- During the 1950s and 1960s, water quality in
form many more experiments, and their combination lakes deteriorated in many areas across North America
of careful long-term observation and experiments has and Europe, with eutrophication being one of the
informed much of our understanding of how forested biggest concerns. A considerable amount of research
catchments work, and in particular how they interact tried to discern the cause. Most of this consisted of
with acid rain. laboratory experiments, often adding nutrients such as
phosphorus or nitrogen, or adding carbon as inorganic
Canada’s Experimental Lakes Area: decades
bicarbonate, to bottles of water from ponds or lakes,
of exploring the consequences of human
and analyzing the response in terms of growth of phy-
activities on lakes
toplankton over a period of a few days to a week. The
Earlier in this chapter, we briefly noted the work by results were conflicting, with many studies suggesting
David Schindler and others on a whole-ecosystem that algal growth and production was primarily lim-
experiment to understand the effects of acid rain on ited by carbon dioxide and bicarbonate availability,
lakes. That experiment was one of many performed at while others indicated control by phosphorus or some-
the Experimental Lakes Area (ELA), a research reserve times nitrogen. Without a clear cause, strong correc-
run by the Canadian government in a sparsely popu- tive policies could not be developed. For instance, if the
lated area of central Canada. ELA was originally estab- problem were phosphorus, one partial solution would
lished in 1968 to study the problem of eutrophication be to ban phosphorus from detergents that contrib-
in lakes: excess growth of algae and cyanobacteria as a uted to wastewater inputs. But without a clear consen-
result of excessive inputs of nutrients. sus, agreement on a phosphorous ban or other policy
 Ecologyandhowtodoit Chapterl mm 21

actions was not possible. If algal growth were really

controlled by the amount of carbon dioxide and bicar-
bonate in a lake, then controlling phosphorus would
be of little value.
In response to concern over eutrophication in the
Great Lakes, the Canadian government established
the Freshwater Institute in 1966, hiring Schindler early
in 1968 to run the whole-lake eutrophication experi-
ment (Figure 1.13). Schindler and his team put in an
access road to the remote area, set up a base camp,
and surveyed many lakes, identifying over 50 that
were suitable for whole-lake experiments. The ELA
group hypothesized that eutrophication was caused by
phosphorus, and they designed a set of experiments to
test this. Their first experiment was a daring one: they
chose a lake (Lake 227) that had extremely low levels
of dissolved carbon dioxide, less than any other lake
anywhere in the world studied to that time, and pro-

Fisheries and Oceans Canada

posed to fertilize it with phosphorus. Lab experiments
with water from Lake 227 had already “proven” that
the low carbon dioxide of the lake strongly limited pro-
duction by algae, but Schindler and his colleagues had
confidence that a whole-lake experiment would yield
a different result. As Schindler (2009) later argued,
if they could prove that phosphorus was actually the FIGURE 1.13 In the Lake 226 experiment, one side of the lake
main stimulant of production even in an ecosystem received nitrogen additions alone, while the other side received both
nitrogen and phosphorus.A plastic barrier prevented exchange of water
such as Lake 227, with extremely low carbon dioxide,
between the two sides of the lake. The stimulation of algal growth by the
it would disprove the carbon hypothesis generally.
phosphorus is clearly seen by the green color in that half of the lake.
Over the winter of 1968-1969, they hauled the Reprinted from Schindler (1977).
phosphorus to the remote lake, using sleds pulled by
snowmobiles. The team began adding phosphorus in
June 1969, and Lake 227 rapidly became eutrophic.
An introduction of an exotic fish species
Despite the low carbon dioxide in the lake, enough
to New Zealand: investigation on
became available to support algal growth, due to dif-
multiple biotic scales
fusion both from the atmosphere and from the car-
bon-rich sediments. The carbon dioxide limitation Historically, studies that encompass more than one or
demonstrated in the laboratory experiments with water two of the four levels in the ecological hierarchy (individ-
from Lake 227 had been an artifact: the rate of supply uals, populations, communities, ecosystems) have been
of new carbon dioxide to the water in the bottles of rare. For most of the 20th century, physiological and
lab experiments was far slower than that to the waters behavioral ecologists (studying individuals), population
of the lake itself. Phosphorus was the real control on dynamicists, and community and ecosystem ecologists
algal growth and eutrophication in lakes. A subsequent tended to follow separate paths, asking different ques-
experiment in another lake divided in half by a plastic tions in different ways. However, over the past decade
barrier showed huge stimulation from fertilization with or two, ecologists have increasingly recognized that,
both phosphorus and nitrogen, compared to nitrogen ultimately, our understanding will be enhanced consid-
alone (look again at Figure 1.13). The publication of erably when the links between all these levels are made
the first results from these experiments led quickly to clear — a point illustrated by studies of an the introduc-
regulation across North America and Europe to reduce tion of an exotic fish to streams in New Zealand.
phosphorus pollution, and the Great Lakes and other Prized for the challenge they provide to anglers,
lakes responded quickly in improved water quality. brown trout (Salmo trutta) have been transported from
The problem of eutrophication in coastal marine eco- their native Europe all around the world for over a cen-
systems has unfortunately taken many decades longer tury. Introduced to New Zealand in 1867, they have
to solve, since the cause there is primarily nitrogen and formed self-sustaining populations in many streams
not phosphorus (Box 1.4). and rivers, often at the apparent expense of native fish
22 Part 1 Introduction

ie)

The deterioration of coastal marine ecosystems

While lake-water quality has improved dramatically as a result of regulations to reduce phosphorus inputs,
coastal marine ecosystems have become steadily more eutrophic since the early 1970s. In fact, by 2007,
a majority of the coastal marine waters in the United States were moderately to severely degraded from
eutrophication, despite the phosphorus controls. Worldwide, harmful algal blooms in coastal waters have
become more common (Figure 1.14), as have areas depleted of oxygen (“dead zones”) such as that in
the northern Gulf of Mexico in the plume of the Mississippi River. Why? Eutrophication of these marine
ecosystems is caused more by nitrogen than by phosphorus, and the regulation of nitrogen pollution has
lagged behind phosphorus regulation by decades in most countries. That nitrogen is the major cause of
coastal eutrophication was well demonstrated in a “mesocosm” experiment on the shore of Narragansett
Bay, Rhode Island, USA. (Mesocosms are experimental arenas large enough to capture many features of
natural systems, in contrast to smaller and less natural ‘microcosms:) Pairs of large fiberglass tanks filled
with seawater from the Bay were enriched with either phosphorus or nitrogen, or left unenriched as con-
trols. The rate of phosphorus input was the same as for the Lake 227 experiment at ELA, described in the
main text. Phosphorus caused no increased in production or amount of algae in the water, but nitrogen
fertilization resulted in much higher quantities and growth of algae (Figure 1.14).
Many factors lead to the differences in the control of eutrophication in lakes and coastal marine eco-
systems. One of the most important is the large supply of phosphorus relative to nitrogen that often enters
coastal marine ecosystems from adjacent oceanic waters. Another factor is nitrogen fixation, the process
whereby bacteria can convert atmospheric molecular nitrogen into biologically available nitrogen. When
nitrogen is in short supply in lakes, cyanobacteria typically increase rapidly in abundance (‘bloom’) and fix

(a)
Gilles Billen, Ph.D

S
Chlorophyll (ug L~)
Robert W. Howarth, Ph.D.

FIGURE 1.14 Massive blooms of harmful algae cause windrows of scum along a beach on the North Sea in the Netherlands
(a, top). Fertilization of 5-meter deep “mesocosms’ filled with seawater from Narragansett Bay (b, bottom) clearly showed a large increase in
algae,as measured by the amount of chlorophyll, from adding nitrogen but not phosphorus (c, right) (Oviatt et al., 1995).
 Ecology and how to doit Chapter 1 a 23

nitrogen, alleviating the shortage and maintaining phosphorus as the limiting nutrient. This response by cya-
nobacteria simply does not occur in most coastal marine ecosystems, as we discuss further in Chapter 11.

| For many years before the mesocosm experiment described above, many short-term lab experiments
with bottles of water from coastal marine ecosystems had shown a response to nitrogen, and not to
phosphorus. Water quality managers tended to discount this evidence, and assume that eutrophication
in coastal waters was the result of the same forces as in lakes. Was this justified?

2 In many cases, the nitrogen that flows to coastal waters and causes eutrophication originates upstream
in river basins at distances of hundreds to a thousand kilometers from the coast, often in different
countries or states. The necessary actions to solve the problems occur upstream, but the damage that
occurs without action is along the coast. How can societies solve coastal eutrophication in an equitable
manner, given this spatial disjunction? Who should pay to reduce the nitrogen pollution?

species such as nonmigratory fish in the genus Galaxias.

When trout are present,
Today, many streams in New Zealand have brown the nymphs are much
In streams without more active at night
trout, many still have only Galaxias, and some have trout, nymphs are when they are less likely
both the introduced and native species of fish. equally obvious to be eaten.
during the day and
Microscopic algae growing on 16 -— at night. , F Day
the individual
rocks compose the base of the food level— BB Night
chain in most south New Zealand consequences for
to

other invertebrate feed-

T

and
Nesameletus visible

streams. Mayfly nymphs

ing behavior
invertebrates graze on the micro-
scopic algae, and they in turn are fed upon by fish.
@

Strikingly, the behavior of nymphs differs depending on

whether they are in Galaxias or brown-trout streams. In
b

one experiment, nymphs collected from a trout stream

and placed in small artificial laboratory channels were
less active during the day than the night, whereas those Galaxias stream Trout stream
collected from a Galaxias stream were active both day Source stream

and night (Figure 1.15). Why might this difference have FIGURE 1.15 Meannumber (+ SE) of Nesameletus ornatus mayfly
evolved? Trout rely principally on vision to capture prey, nymphs collected either from a trout stream or a Galaxias stream that were
whereas Galaxias rely on mechanical cues. Thus, inver- recorded by means of video as visible on the substrate surface in labora-
tebrates in a trout stream are considerably more at risk tory stream channels during the day and night (in the absence of fish).
of predation during daylight hours when the trout can Mayflies from the trout stream are more nocturnal than their counterparts
see best and are most active. from the Galaxias stream. (After Mcintosh & Townsend, 1994.)
That an exotic . predator such the community -
as brown trout has direct effects on brown trout cause fish. After a further 12 days, invertebrates and algae
mayfly behavior is not surprising. a cascade
of were sampled (Figure 1.16). A significant effect of
effects
Are there broader effects that cas- brown trout reducing invertebrate biomass was evident
cade through the community to other species? An (P = 0.026), but the presence of Galaxias had no effect
experiment involving artificial flow-through channels on invertebrate biomass compared to the no-fish con-
(several meters long, with mesh ends to prevent escape trol. Algal biomass achieved its highest values, too, in
of fish but to allow invertebrates to colonize naturally) the trout treatment (P = 0.02), probably because there
placed in a stream tested whether the exotic trout affect were fewer invertebrates grazing the algae.
the stream food web differently from the displaced What are the consequences
. the ecosystem —
native Galaxias. Three treatments were established: no to the ecosystem, in terms of over- trout and energy
fish, Galaxias present, and trout present, at naturally all photosynthesis by the algae flow
occurring densities. Algae and invertebrates were and energy flow to the mayflies and
allowed to colonize for 12 days before introducing the other invertebrates, and ultimately to the fish? This
24 8 Part Introduction

was investigated in two neighboring tributaries the trout stream than in the Galaxias stream. This
of the Taieri River, one where brown trout was the resulted in more energy to support the entire food
only fish species and one containing only Galaxias. web, and the invertebrates that eat algae produced
Photosynthesis by the algae was six times greater in new biomass in the trout stream at about 1.5 times the
rate found in the Galaxias stream. Trout themselves
produced new biomass at roughly nine times the rate
that Galaxias did (Figure 1.17). Thus, the algae, inver-
tebrates and fish are all more productive in the trout
4 3
stream, than in the Galaxias stream.
Ca Does this mean that the introduction of the non-
E re native species, brown trout, was beneficial for New
2s E
o 2 Zealand streams? Yes, if the sole goal is to have a high
g z
53 level of fish production, and keep anglers happy. Of
2 g
© € course, invasive species — whether introduced on purpose
ga $= 1 or otherwise — are a huge and increasing problem, and
‘a 2
2 @
very often have a net negative influence on both basic
ecological function and on a variety of issues directly
connected to human well-being. This is a major topic
. 0 gy
N G a for ecologists, and is explored further later in the text.
m.. predation =e Fish predation regime

FIGURE 1.16 (a) Total invertebrate biomass and (b) algal bio- Why Asian vultures were heading for
mass (chlorophyll a) (+ SE) for an experiment performed in summer extinction: The value of a modeling study
in a small New Zealand stream. In experimental replicates where trout
are present, grazing invertebrates are rarer and graze less; thus, algal In 1997, vultures in India and Pakistan began drop-
biomass is highest. G, Galaxias present; N, no fish;T, trout present. (After ping from their perches. Local people were quick to
Flecker & Townsend, 1994.) notice dramatic declines in numbers of the oriental

9 Production
Demand
Production/demand (g AFDW m~)

150

Galaxias Trout Galaxias Trout Galaxias Trout

Algae Invertebrates Fish

FIGURE 1.17 Annual estimates for ‘production’ of biomass at one trophic level, and the ‘demand’
for that biomass (the amount consumed)
at the next trophic level, for (a) primary producers (algae), (b) invertebrates (which consume algae), and (c) fish (which consume invertebrates).
Estimates are for a trout stream and a Galaxias stream. In the former, production at all trophic levels is higher, but because the trout consume
essentially all of the annual invertebrate production (b), the invertebrates consume only 21% of primary production (a). In the Galaxias stream,
these fish consume only 18% of invertebrate production, ‘allowing’ the invertebrates to consume the majority (75%) of annual primary production.
AFDW stands for ash-free dry weight. (After Huryn, 1998.)
 Ecologyandhowtodoit Chapter] mm 25

2 =N,/N,

Baseline Baseline
survival, survival,
Ss Ms, ff s
. Maturation
aural and survival
FRectof ~~ Elect of
diclofenac diclofenac

> Drpape Rate at which Erobapalty Rate at which

E ofa carcass ofa carcass
= containing ees containing oe
3 diclofenac, oie diclofenac, ape
3
a ¢ .
FIGURE 1.18 Flow diagram showing the elements of a model of how the number of adult vultures in the population changes from one
year (N,_,) fo the next (N,). The oriental white-backed vulture, whose populations have shown disastrous declines in India and Pakistan, is shown in
the inset. The number of adult vultures in year tf depends on the number present the previous year (f —1), some of which die from natural causes
(baseline survival) and others because of diclofenac poisoning. The number of adults in year f also depends on the number of vultures born 5 years
previously (f— 5), because vultures do not mature until they are 5 years old. Again, some newborn vultures die before maturity from natural causes
and others because of diclofenac poisoning. The reduction in survival due to diclofenac depends on two things: the probability that a carcass contains
diclofenac (C) and the rate at which carcasses are eaten (F). (© Alamy Images AGJX38.)

white-backed vulture Gyps bengalensis (Figure 1.18) that carcasses of domestic animals treated with diclof-
and the long-billed vulture G. indicus. Repeated popu- enac were lethal to captive vultures. Diclofenac, a non-
lation surveys from 2000 to 2003 confirmed alarming steroidal anti-inflammatory drug developed for human
rates of decline: between 22% and 50% of the popula- use in the 1970s, had only recently come into common
tion was being lost each year. This was of very great use as a veterinary medicine in Pakistan and India.
concern because of the crucial role vultures play in Thus, a drug that benefited domestic mammals proved
everyday life for people in India and Pakistan, dispos- lethal to the vultures that fed on their bodies.
ing of the dead bodies of large animals, both wild and The circumstantial evidence
domestic. The loss of vultures enhanced carrion avail- was strong, but given the rela-
ability to wild dogs and rats, allowing their popula- tively small numbers of diclofenac-
tions to increase and raising the probability of diseases contaminated dead bodies available
such as rabies and plague being transmitted to humans. to wild vultures, was the associated vulture mortality suf-
Moreover, contamination of nearby wells and the spread ficient explanation for the population crashes? Or might
of disease by flies became more likely now that dead other factors also be at play? Were there other toxins?
animals were not quickly picked clean by vultures. One This was the question addressed by Rhys Green and his
group of people, the Parsees, were even more intimately team by means of a simulation population model (Green
affected, because their religion calls for the dead to be et al. 2004). On the basis of their surveys of popula-
taken in daylight to a special tower (dakhma) where the tion declines and knowledge of birth, death and feeding
body is stripped clean by vultures within a few hours. It rates, the researchers built a mathematical model to pre-
was crucial for ecologists to quickly determine the cause dict the behavior of the vulture populations. We show
of vulture declines so that action could be taken. their model as a flow diagram (Figure 1.18).
It took scientists a few years to find a common The model was based on a series of simple and
element in the deaths of otherwise healthy birds — each well-articulated assumptions. For example, they
had suffered from visceral gout (accumulation of uric assumed that vultures do not breed until they are five
acid in the body cavity) followed by kidney failure. years old, and then are capable of rearing only one
Soon a crucial piece in the jigsaw became clear: vultures juvenile per year, but only if both parents survive the
dying of visceral gout contained residues of the drug breeding season of 160 days. Other model inputs spec-
diclofenac (Oaks et al., 2004). Then it was confirmed ify normal birth and death rates for the population,
26 = Part1_ Introduction

but also how survival may depend on dicofenac poi- 1 Models can be valuable for exploring scenarios and
soning. This reflects the probability an adult will eat situations for which we do not have, and perhaps
from a diclofenac-affected carcass, which in turn cannot expect to obtain, real data (e.g., what would
depends partly on the proportion of carcasses in the be the consequences of different baseline survival or
environment that contain diclofenac and partly on feeding rates?).
how often vultures feed. As you might guess, there is a
2 They can be valuable, too, for summarizing our cur-
lot of complexity, and a lot of good biology, behind
rent state of knowledge and generating predictions
these assumptions.
in which the connection between current knowledge,
Mathematical formulae were
assumptions, and predictions is explicit and clear
developed to predict changes in
(given various values for S and F in Figure 1.18, and
population size, but the details
knowing A, the change in population from one year
need not concern us here. More
to the next, what values of C do these imply?).
specifically, the researchers posed
the question: what proportion of carcasses (C) would 3 In order to be valuable in these ways, a model does
have to contain lethal doses of diclofenac to cause the not have to be (indeed, cannot possibly be) a full
observed population declines? The model showed that and perfect description of the real world it seeks to
only 1 in 135 carcasses would have to contain diclofe- mimic — all models incorporate approximations (the
nac to cause the observed population decline. The pro- vulture model was, of course, a very ‘stripped down’
portions of vultures found dead or dying in the wild version of its true life history).
with signs of diclofenac poisoning were closely similar
4 Caution is therefore always necessary — all conclu-
to the proportions of deaths expected from the model sions and predictions are provisional and can be
if the observed population decline was due entirely
no better than the knowledge and assumptions on
to diclofenac poisoning. The researchers concluded,
which they are based - but applied cautiously they
therefore, that diclofenac poisoning was a sufficient
can be useful (the vulture model prompted changes
cause for the dramatic decline of wild vultures. As a
in management practices and research into new
result of this research, governments have taken action
drugs).
to ban the use of diclofenac.
This example, then, has illustrated a number of 5 Nonetheless, a model is inevitably applied with
important general points about mathematical models much more confidence once it has received support
in ecology: from real sets of data.

The science of ecology Ecology in practice

Ecology is the scientific study of the distribution and abundance Ecologists use a wide variety of approaches including com-
of organisms, the interactions that determine that distribution parative observations, experiments, and models. Experiments
and abundance, and the relationships between organisms and include natural experiments such as the observation of agri-
the transformation and flux of energy and matter. cultural fields that were abandoned at various times over the
past century, small scale experiments such as the addition of
Scales, diversity of approaches, and rigor nitrogen to small plots, and whole-ecosystem experiments such
Ecology deals with four levels of ecological organization: indi- as cutting all the trees from a catchment or adding phosphorus
vidual organisms, populations, communities, and ecosystems. to a whole lake.
Ecology is a rigorous science using standard scientific techniques.
 Ecologyandhowtodoit Chapter] mM 27

Why do some temporal patterns in ecology need long runs When all the trees were felled in a Hubbard Brook catch-
of data to detect them, while other patterns need only short ment, there were dramatic differences in the chemistry of
runs of data? the stream water draining the catchment. How do you think
stream chemistry would change in subsequent years as
Discuss the pros and cons of descriptive studies as opposed
plants begin to grow again in the catchment area?
to experiments of the same ecological phenomenon.
Why should we have confidence in whole-lake experi-
What is a ‘natural field experiment’? Why are ecologists
ments, even though they are frequently performed without
keen to take advantage of them?
replication?
Search the library for a variety of definitions of ecology;
By what mechanism did the introduction of brown trout in
which do you think is most appropriate and why?
New Zealand streams lead to changes in the production of
How might the results of the Cedar Creek study of old-field algae?
succession have been different if a single field had been
What are the main factors affecting the confidence we can
monitored for 50 years, rather than simultaneously compar-
have in predictions of a mathematical model?
ing fields abandoned at different times in the past?

Challenge Questions
] Discuss the different ways that ecological evidence can be The variety of microorganisms that live on your teeth have
gained. How would you go about trying to answer one of an ecology like any other community. What do you think
ecology’s unanswered questions, namely, ‘Why are there might be the similarities in the forces determining species
more species in the tropics than at the poles?’ richness (the number of species present) in your oral com-
munity as opposed to a European grassland?
 BI
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Chapter

Ecology's Evolutionary Backdrop

CHAPTER CONTENTS
2.1 Evolution by natural selection 2.5 Continental drift, parallel and convergent
2.2 Evolution within species evolution
2.3. The ecology of speciation 2.6 Conclusion
2.4 The effects of climatic change on the
evolution and distribution of species

KEY CONCEPTS

After reading this chapter you will be able to:

* explain that Darwin and Wallace, who were explain that the evolutionary history of species
responsible for the theory of evolution by natural constrains what future selection can achieve
selection, were both, essentially, ecologists
compare and contrast convergent and parallel
* appreciate that natural selection can act very evolution
quickly on heritable variation, and that we can
explain that natural selection fits organisms to
study it in action and control it in experiments
their past, rather than anticipating their future
° identify what is required for the origin of new
species
 Ecology's Evolutionary Backdrop ~Chapter2 @ 29

\ s the great Russian-American biologist Dobzhansky said, “Nothing in biology

makes sense, except in the light of evolution.” But equally, very little in
evolution makes sense except in the light of ecology: ecology provides the
stage directions through which the “evolutionary play” is performed. Ecologists and
evolutionary biologists need a thorough understanding of each other's disciplines
to make sense of key patterns and processes.

2.1 EVOLUTION BY NATURAL requires an understanding of the processes of evolution

SELECTION that have led to present-day patterns of diversity and
distribution.
The Earth is inhabited by very many different types Until relatively recently, the »
of organism. They are distributed neither randomly | emphasis with biological diversity
nor evenly over the surface of the globe. Any sampled was on using it (for example, for
area contains only a small subset of the variety of | medicine), exhibiting it in zoos and
_ almost everywhere
species present on Earth. Why are there so many types _ botanic gardens, or cataloging it in
of organism? Why are the distributions of so many museums (Box 2.1).Yet such catalogs are more like
so restricted? Answering these ecological questions stamp collecting than science unless accompanied by

2.1. Historical Landmarks

A Brief History of the Study of Diversity

An awareness of the diversity of living organisms, and of what lives where, is part of the knowledge that the
human species accumulates and hands down through the generations. Hunter—-gatherers needed (and
still need) detailed knowledge of the natural history of the plants and animals in their immediate environ-
ment in order to obtain food successfully and escape the hazards of being poisoned or eaten.
More than 4,000 years ago, the Chinese emperor Shen Nung compiled what was perhaps the first
written ‘herbal’ of useful plants; and by around 2,000 years ago, the Greek, Dioscorides, had described 500
species of medicinal plants and illustrated many of them in his De Materia Medica, which remained in use
through the following 1,500 years.
Collections of living specimens in zoos and botanic gardens also have a long history — certainly
back to Greece almost 3,000 years ago. Then, in the 17th century, the urge to collect from the diversity
of nature developed in the West, allowing some individuals to make their living by finding interesting
specimens for other people's collections. For example, John Tradescant the father (died 1638) and John
Tradescant the son (1608-1662) spent most of their lives collecting plants abroad for the gardens of the
British aristocracy. The father was the first English botanist to visit Russia (1618), bringing back many living
plants. His son made three visits to collect specimens in the American colonies.
Wealthy individuals built up vast collections into personal museums and traveled or sent travelers in
search of novelties from new lands as they were discovered and colonized. Naturalists and artists (offen the
same people) were sent to accompany the major voyages of exploration, to report and take home, dead or
alive, collections of the diversity of organisms and artifacts that they found. Taxonomy (giving names to the
various types of organism) and systematics (organizing and classifying them) developed and flourished.
Random documents with unrelated
content Scribd suggests to you:
tons, was launched in December of 1907, and is one of the most
notable productions of recent years. She is spar-decked throughout,
with magnificent lines and a handsome appearance, whilst retaining
the more conventional stem-plus-bowsprit. She has exceptional
accommodation, all connected by corridors and vestibules with no
fewer than a dozen state-rooms for guests. She is driven by two sets
of triple-expansion engines actuating twin-screws, which, to minimise
vibration, are at a different pitch, and run at varying speeds. She can
carry sufficient coal to allow her to cruise for 6,000 miles, and both in
internal and external appearance is as handsome as she is capable.

THE S.Y. “SAGITTA.”

From a Photograph. By permission of Messrs. Camper & Nicholson, Ltd.
 THE S.Y. “TRIAD.”
From a Photograph. By permission of the Caledon Shipbuilding Co., Ltd.

With the capabilities of which the motor has shown itself to be

possessed, the future of the steam yacht is perhaps a little uncertain.
Economy would seem to indicate that the former has numerous
merits in that it enables sail power to be utilised more readily, and
thus may arrest the fashion which is advancing in the direction of
steam. For long passages the extreme comfort which is now
obtainable in the modern liner leaves no choice in the matter. To
keep up a steam yacht for the usual summer season of four months
is a very serious item of expenditure. If we reckon £10 per ton as the
average cost—and this is the accepted estimate—it will be seen that
such a yacht as the Wakiva, for instance, leaves but little change out
of £10,000 per year, and for this expenditure most men would expect
to get a very large return in the way of sport and travel. Whether or
not a like proportionate return is made, at least in giving employment
to thousands of shipbuilding and yacht-hands, this special branch of
sea sport is deserving of the high interest with which it is regarded.
 CHAPTER XI
THE BUILDING OF THE STEAMSHIP

We propose in the present chapter, now that we have seen the

evolution of the steamship through all its various vicissitudes and in
its special ways, to set forth within the limited space that is now left
to us some general idea of the means adopted to create the great
steamship from a mass of material into a sentient, moving being.
Around the building of a ship there is encircling it perhaps far
more sentiment than in the activity of almost any other industry.
Poets and painters have found in this a theme for their imagination
not once, but many times. Making a ship is something less prosaic, a
million times more romantic, than making a house, for the reason
that whilst the ship, as long as she remains on the stocks, is just so
many thousand tons of material, yet from the very moment when she
first kisses the water she becomes a living thing, intelligent, with a
character of her own, distinct and recognisable. In the whole
category of man-made things there is nothing comparable to this.

Fig. 1.—FLUSH-DECKED TYPE.

 Fig. 2.—“THREE ISLAND,” TYPE.

Fig. 3.—TOP-GALLANT FORECASTLE TYPE.

Fig. 4.—TOP-GALLANT FORECASTLE TYPE, WITH

RAISED QUARTER-DECK.

Fig. 5.—EARLY “WELL-DECK” TYPE.

Her genesis begins when the future owners resolve to have her
built. Before any plans are drawn out there must first be decided the
dimensions, the displacement and the general features which she is
to possess, whether she is to be a slow ship, a fast ship, engaged in
passenger work, cargo-carrying, on the North Atlantic route, for the
East through the Suez Canal, and so on; for all these factors
combine to determine the lines on which she is to be built. Before we
progress any farther, let us get into our minds the nine different types
which separate the generic class of steamships. If the reader will
follow the accompanying illustrations, we shall not run the risk of
being obscure in our argument. Fig. 1, shows the steamship in its
elementary form, just a flush-decked craft, with casings for the
protection of the engines as explained on an earlier page. This
represents the type of which the coasting steamer illustrated
opposite page 134 is an example. This casing in the diagram before
us is, so to speak, an island on the deck, but presently it was so
developed that it extended to the sides of the ship, and, rising up as
a continuation of the hull, became a bridge. At the same time a
monkey forecastle and a short poop were added to make her the
better protected against the seas. This will be seen in Fig. 2. This is
known as the “three-island” type for obvious reasons. It must be
understood that on either side a passage leads beneath the bridge-
deck so as to allow the crew to get about the ship. But from being
merely a protection for the bows of the ship, the monkey forecastle
became several feet higher, so that it could accommodate the
quarters of the crew, and this “top-gallant” forecastle, as it is known,
will be seen in Fig. 3. At the same time, the short poop or hood at the
stern has now become lengthened into something longer. But in Fig.
4 we find the lengthened poop becoming a raised quarter-deck—that
is, not a mere structure raised over the deck, but literally a deck
raised at the quarter. This raised quarter-deck was the better able to
withstand the violent force of the sea when it broke over the ship. In
Fig. 5 we have a still further development in which the topgallant
forecastle is retained as before, but the long poop and the after end
of the bridge are lengthened until they meet and form one long
combination. This is one of the “well-deck” types, the “well” being
between the after end of the forecastle and the forward end of the
bridge-deck. This well was left for the reason that it was not required
for carrying cargo, because it was not desirable to load the ship
forward lest she might be down at the head (which in itself would be
bad), whilst at the same time it would raise the stern so that the
propeller was the more likely to race. But in the modern evolution of
the steamship it is not only a question of trim and seaworthiness that
have been taken into consideration, but also there are the rules and
regulations which have been made with regard to the steam vessel.
Now, this well-space not being reckoned in the tonnage of the ship
(on which she has to pay costly dues) if kept open, it was good and
serviceable in another way. Considered from the view of
seaworthiness, this well, it was claimed, would allow the prevention
of the sweeping of the whole length of the ship by whatever water
that broke aboard the bows (which would be the case if the well were
covered up). If left open, the water could easily be allowed to run out
through the scuppers. But this type in Fig. 5 is rather midway in the
transition between the “three-island” type and the shelter-deck type.
The diagram in Fig. 6 is more truly a well-decker, and differs from the
ship in Fig. 5, in that the one we are now considering has a raised
quarter-deck instead of a poop. She has a top-gallant forecastle, a
raised quarter-deck and bridge combined, and this type was largely
used in the cargo ships employed in crossing the Atlantic Ocean. It is
now especially popular in ships engaged in the coal trade. The
advantages of this raised quarter-deck are that it increases the cubic
capacity of the ship, and makes up for the space wasted by the shaft
tunnel. By enabling more cargo to be placed aft, it takes away the
chance of the ship being trimmed by the head.

Fig. 6.—“WELL-DECK” TYPE.

 Fig. 7.—“SPAR-DECK” TYPE.

Fig. 8.—“AWNING-DECK” TYPE.

Fig. 9.—“SHADE-DECK” TYPE.

Fig. 7 shows a “spar-decker,” which is the first of the three-

deckers that we shall now mention. This was evolved for the purpose
of carrying passengers between decks. It has a continuous upper
deck of fairly heavy construction, the bridge deck, of course, being
above the spar deck. In Fig. 8 we have the “awning-decker,” which
has a continuous deck lighter in character than the last-mentioned
type, and like the latter, the sides are completely enclosed above the
main deck. Because of this lightness of construction, it is not
customary to add further erections above that are of any weight. Its
origin was due to the desire to provide a shelter for the ships
employed in carrying Oriental pilgrims. Later on this type was
retained in cargo-carriers. Finally, we have the “shade-decker” as in
Fig. 9, which is provided with openings at the side for ventilation.
This type is so well known to the reader from posters and
photographs, that it is scarcely essential to say much. But we may
remark that the lightly constructed deck fitted between the poop and
forecastle is supported by round stanchions, open at the sides (as
shown herewith), but sometimes closed by light plates. It is built just
of sufficient strength to provide a promenade for passengers, or
shelter for cattle, on the upper deck. This is still a very popular type
for intermediate and large cargo steamers.

THE BUILDING OF THE “MAURETANIA.”

Showing Floor and part of Frames.
From a Photograph. By permission of the Cunard Steamship Co.

With these different types before us, we may now go on with our
main subject. Having settled the question as to the type and
character of the steamship to be built, the next thing is to design the
midship section, which shows the general structural arrangements
and scantlings of the various parts. In the drawing-office the plans
are prepared, and the various sections of the ship worked out by
expert draughtsmen attached to the shipbuilding yard. This
necessitates the very greatest accuracy, and the building is usually
specially guarded against those who might like to have an
opportunity of obtaining valuable secrets. The plans having been
worked out on paper, there follows the “laying off” on the floor of an
immense loft, called the “mould floor,” where the plans are
transferred according to the exact dimensions that are to be
embodied in the ship. In many cases the future owner insists on a
wooden model being submitted in the first instance, by the builder,
so that a fair idea may be obtained of the hull of the proposed ship.
Each vessel is known at the shipbuilder’s by a number and not
by her name. The keel is the first part of her to be laid, which
consists of heavy bars of iron laid on to blocks of wood called
“stocks,” and the line of these slants gently down to the water’s
edge, so that when, after many months, the time arrives for the
launching of the great ship, she may slide down easily into the sea
that is, for the future, to be her support. After these bars have been
fastened together, then the frames or ribs are erected, the ship being
built with her stern nearest to the water, and her bow inland, except
in the few cases (as, for example, that of the Great Eastern), where
a vessel, owing to her length in proportion to the width of the water-
space available, has to be launched sideways. These ribs are bent
pieces of steel, which have been specially curved according to the
pattern already worked out. Let us now turn to the accompanying
illustrations which show the steamship in course of construction.
These have been specially selected in order that the reader might be
able to have before him only those which are of recent date, and
show ships whose names, at least, are familiar to him.
 THE “GEORGE WASHINGTON” IN COURSE OF CONSTRUCTION.
Showing Framing from the Stern.
From a Photograph. By permission of the Norddeutscher Lloyd Co.

The photograph opposite page 286 represents the Mauretania

being built on the Tyne. This striking photograph shows the floor and
the double cellular bottom of the leviathan in the foreground; whilst in
the background the frames of the ship have been already set up.
Some idea of the enormous proportions may be obtained from the
smallness of the men even in the foreground. The next illustration
represents the Norddeutscher Lloyd liner, George Washington, and
exhibits the framing of the ship and bulkheads before the steel-
plating had been put on. The photograph was taken from the stern,
looking forward, and one can see already the “bulge” which is left on
either side to allow for the propeller shafts. Opposite page 290 is
shown the bow end of the Berlin (belonging to the same company) in
frame, and on examining her starboard side it will be seen that
already some of her lower plates have been affixed. Finally, opposite
page 292 is shown one of the two mammoth White Star liners in
course of construction. This picture represents the stern frame of the
Titanic as it appeared on February 9th, 1910. No one can look at
these pictures without being interested in the numerous overhead
cranes, gantries and scaffolding which have to be employed in the
building of the ship. The gantries, for instance, now being used at
Harland and Wolff’s Belfast yard are much larger than were used
even for the Celtic and Cedric, and have electric cranes, for handling
weights at any part of the berths where the ships are being built.
Cantilever and other enormous cranes are also employed. Cranes
are also now used in Germany fitted with very strong electro-
magnets which hold the plates by the power of their attraction, and
contribute considerably to the saving of labour.
Whilst the hull of the ship is being built, the engines are being
made and put together in the erecting-shop—which also must needs
have its powerful cranes—and after being duly tested, the various
parts of the engines are taken to pieces again and erected
eventually in the ship after she has been launched. After the frames
and beams are “faired” the deck-plating is got in hand. Besides
affording many advantages, such as promenades and supports for
state-rooms, the deck of a ship is like the top of a box, and gives
additional strength to a ship. The illustration opposite page 292
shows the shelter deck of the Orient liner Orsova. The photograph
was taken looking aft, on August 1st, 1908, whilst the ship was being
built at Messrs. John Brown & Co.’s yard, Clydebank. The
photograph is especially interesting as showing the enormous
amount of material which has to go to the making of the steamship.
But even still more significant is the next illustration, which shows
one of the decks of the Lusitania whilst in course of construction. To
the average man it seems to be well-nigh impossible ever to get
such masses into the water.
BOWS OF THE “BERLIN” IN COURSE OF
CONSTRUCTION.
From a Photograph. By permission of the Norddeutscher
Lloyd Co.
 THE “BERLIN” JUST BEFORE HER
LAUNCH.
From a Photograph. By permission of the
Norddeutscher Lloyd Co.

After the plates have been all fastened by rivets to the frames,
and the outside of the ship has been given a paint of conventional
salmon pink, the time approaches for her to be launched. During her
building the ship has been resting on the keel blocks where her
centre touches, but her bilges have been supported by blocks and
shores. These latter will be seen in the illustration of the Mauretania
already considered. As the day for launching approaches, so also
does the anxiety of the builders increase, for at no time in her career
is the ship so seriously endangered. On the day of the launch the
weight of the vessel is gradually transferred from the stocks on which
she has been built, to the cradle, being lifted bodily from the keel-
blocks by means of an army of men driving wedges underneath her
bottom. This cradle is constructed on the launching ways, and the
ship herself, being now “cradle-borne,” is held in place only by a
number of props called “dog-shores.” At the right moment the signal
is given for these to be knocked aside, and at the first symptoms of
the ship in her cradle showing an inclination to glide, the bottle of
wine is broken against her bows by the lady entrusted with so
pleasant an honour. With a deep roar the ship goes down the ways,
and as soon as the vessel becomes waterborne the cradle floats.
The ship herself is taken in charge by a tug, whilst numerous small
boats collect the various pieces of timber which are scattered over
the surface of the water. Two or three days before the launch, the
cradle which has been fitted temporarily in place, is taken away and
smeared with Russian tallow and soft soap. The ways themselves
are covered with this preparation after they have been well scraped
clean. In case, however, the ship should fail to start at the critical
moment after the dog-shores have been removed, it is usual now to
have a hydraulic starting ram (worked by a hand-pump) under the
forefoot of the ship. This will give a push sufficiently powerful to start
the great creature down her short, perilous journey into the world of
water which is to be her future abiding-place.
But it can readily be imagined that such a ponderous weight as
this carries a good deal of impetus with it, and since in most cases
the width of the water is confined, precautions have to be taken to
prevent the ship running ashore the other side and doing damage to
herself—perhaps smashing her rudder and propellers, or worse.
Therefore, heavy anchors have been buried deep into the ground,
and cables or hawsers are led from the bows and quarters and
attached thereto, or else to heavy-weights composed of coils of
chain, whose friction over the ground gradually stops the vessel. Not
infrequently the cables break through the sudden jerk which the
great ship puts on them, and the anchors tear up the slip-way.
Perhaps as many as eight cables may be thus employed, each being
made fast to two or three separate masses of about five to fifteen
tons, but with slack chain between so that only one at a time is
started. As soon as the ship has left the ways, all the cables become
taut, and they put in motion the first lot of drags. Further on, the next
lot of drags receive their strain, then the third, so that no serious jerk
may have been given, and the ship gradually brings up owing to the
powerful friction. Lest the force of the ship going into the water
should damage the rudder or the propeller, these, if they have been
placed in position, are locked so as to prevent free play. After this the
ship is towed round to another part of the yard where her engines
are slung into her by means of powerful cranes. The upper
structures are completed, masts stepped and an army of men work
away to get her ready for her builders’ trials. Carpenters are busy
erecting her cabins, painters and decorators enliven her internal
appearance, and upholsterers add the final touches of luxury to her
saloons and lounges.
 STERN FRAME OF THE “TITANIC,” FEB. 9, 1910.
From a Photograph. By permission of Messrs. Ismay, Imrie & Co.

Turning now to the illustration facing page 290, we see the

Norddeutscher Lloyd Berlin just before she was launched. The
anchors and cables which will be dropped as soon as she has
floated will be seen along her port side, and the platform for her
christening is already in place. In the illustration facing page 294,
which shows the launch of the Royal Mail Steam Packet Company’s
Araguaya, we have a good view afforded of the ship as she is just
leaving the ways and becoming water-borne. The other illustration
on the same page shows the launch of one of those turret-ships to
which reference was made in an earlier chapter. In the picture of the
Berlin will be seen the system of arranging the steel plates in the
construction of the ship, and the rivets which hold them in place.

THE SHELTER DECK OF THE “ORSOVA” IN COURSE OF

CONSTRUCTION.
From a Photograph. By permission of Messrs. Anderson, Anderson & Co.
 ONE OF THE DECKS OF THE “LUSITANIA” IN COURSE
OF CONSTRUCTION.
From a Photograph. By permission of the Cunard Steamship Co.

One of the most important events of the ship’s life is her trial trip.
Before this occurs the ship’s bottom must be cleaned, for a foul
underwater skin will deaden the speed, and give altogether
erroneous data. The weather should be favourable also, the sea
calm, and the water not too shallow to cause resistance to ships of
high speed, while a good steersman must be at the helm so as to
keep the ship on a perfectly straight course. Around our coasts at
various localities are noticeable posts erected in the ground to
indicate the measured mile. To obtain the correct data as to the
speed of the ship, she may be given successive runs in opposite
directions over this measured mile; a continuous run at sea, the
number of revolutions being counted during that period, and a
continuous run past a series of stations of known distances apart,
the times at which these are passed being recorded as the ship is
abreast with them. For obtaining a “mean” speed over the measured
mile, one run with the tide and one against the tide supply what is
required. During these trials, the displacement and trim of the ship
should be as nearly as possible those for which she has been
designed. But besides affording the data which can only show
whether or not the ship comes up to her contract, these trials are
highly valuable as affording information to the builder for subsequent
use, in regard both to the design of the ship herself and the amount
of horsepower essential for sending her along at a required speed.
The amount of coal consumption required is also an important item
that is discovered. This is found as follows: Let there be used two
bunkers. The first one is not to be sealed, but the latter is. The
former is to be drawn upon for getting up steam, taking the ship out
of the harbour, and generally until such time as she enters upon her
trial proper. This first bunker is then sealed up, and the other one
unsealed, and its contents alone used during the trial. After the trial
is ended, the fires being left in ordinary condition, the second bunker
is again sealed up, and the first bunker drawn upon. By reckoning up
the separate amounts it is quite easy afterwards to determine the
exact quantity which the ship has consumed during a given number
of knots in a given time. Finally, after every detail has been
completed, the ship is handed over to her owners and steams away
from the neighbourhood of her birth. Presently she arrives at her
port, whence she will run for the next ten or twenty years, and before
long she sets forth with her first load of passengers, mails and cargo
on her maiden trip across the ocean. To begin with, she may not
establish any new records for speed; for a ship takes time to find
herself, and her officers to understand her individualities. “Know your
ship” is one of the mottoes which an ambitious officer keeps ever
before him, and if this is true on the navigation bridge, it is even still
more true down below, where the engines will not show their full
capabilities for several passages at least.
 LAUNCH OF THE “ARAGUAYA.”
From a Photograph. By permission of the Royal Mail Steam Packet Co.

LAUNCH OF A TURRET-SHIP.
From a Photograph. By permission of Messrs. Doxford & Sons, Sunderland.
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