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Ecology, Second Edition
by Bill Freedman, Jeffery A. Hutchings, Darryl T. Gwynne, John P. Smol, Roger Suffling,
Roy Turkington, Richard L. Walker, and Dawn Bazely
Vice President, Editorial Production Service: Cover Design:

Higher Education: Integra Software Services Pvt. Ltd. Jennifer Leung
Anne Williams
Copy Editor: Cover Image:
Publisher: Karen Rolfe Paul Nicklen/National Geographic
Paul Fam Images
Proofreader:
Marketing Manager: Integra Software Services Pvt. Ltd. Box Images:
Leanne Newell Ecology in Depth, Thinkstock;
Indexer: Environmental Applications,
Developmental Editor: Integra Software Services Pvt. Ltd. Evlakhov Valeri/Shutterstock;
Toni Chahley A Canadian Ecologist,
Design Director:
© iStockphoto.com/Bart Coenders
Photo Researcher/Permissions Ken Phipps
Coordinator: Compositor:
Kristiina Paul Managing Designer:
Integra Software Services Pvt. Ltd.
Franca Amore
Senior Production Project
Manager: Interior Design:
Imoinda Romain Dianna Little
COPYRIGHT © 2015, 2011 Every effort has been made to Revision of: Ecology : a Canadian
by Nelson Education Ltd. trace ownership of all copyrighted context / Bill Freedman ... [et al.].
material and to secure permission — Toronto : Nelson Education,
WCN: 02-200-201 from copyright holders. In the [2010], c2011.
event of any question arising as Includes bibliographical references
Printed and bound in the United
to the use of any material, we will and index.
States of America
be pleased to make the necessary ISBN 978-0-17-651014-5 (bound)
1 2 3 4 17 16 15 14
corrections in future printings.
1. Ecology—Canada—Textbooks.
For more information contact
Library and Archives Canada I. Hutchings, Jeffrey Alexander,
Nelson Education Ltd.,
Cataloguing in Publication 1958-, author II. Gwynne, Darryl
1120 Birchmount Road, Toronto,
T., author III. Smol, John Paul,
Ontario, M1K 5G4. Or you can visit Freedman, Bill, author 1955-, author IV. Suffling, Roger,
our Internet site at Ecology : a Canadian context / 1948-, author V. Turkington,
http://www.nelson.com Bill Freedman, Department of Roy, 1951-, author VI. Walker,
Biology, Dalhousie University, Richard L. (Professor of biology),
ALL RIGHTS RESERVED. No part of
Jeffrey A. Hutchings, Department author VII. Bazely, Dawn, author
this work covered by the copyright
of Biology, Dalhousie University, VIII. Title.
herein may be reproduced,
Darryl T. Gwynne University of
transcribed, or used in any form or
Toronto in Mississauga, John P. QH541.E355 2014
by any means—graphic, electronic,
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or mechanical, including
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photocopying, recording, taping, ISBN-13: 978-0-17-651014-5
Faculty of Environment, University
Web distribution, or information ISBN-10: 0-17-651014-1
of Waterloo, Roy Turkington
storage and retrieval systems—
Department of Botany, and Beaty
without the written permission of
Biodiversity Research Centre,
the publisher.
University of British Columbia,
For permission to use material Richard L. Walker, Department
from this text or product, submit of Biological Sciences, University
all requests online at of Calgary, Dawn Bazely, Biology
www.cengage.com/permissions. Department, York University.
Further questions about — Second edition.
permissions can be emailed to
permissionrequest@cengage.com but when there is a seasonal run upper reaches of the watersheds
of Pacific salmon (Oncorhynchus) of the streams where they breed.
About the Cover Image: This they focus on those fish as a highly Feeding by the bears extends this
image shows a Kermode bear nutritious and energy-rich food. The ecosystem function by transferring
(Ursus americanus kermodei) in whitish fur is due to recessive genes nutrients in the form of partially
its habitat of the Pacific coastal and is not an albino trait (the bears eaten fish carcasses and defecations
forest of western and northern do not have red eyes). The light to terrestrial parts of the watershed.
British Columbia and the Alaskan colour may be less visible to salmon The whitish Kermode bears are
panhandle. The Kermode bear and so may confer a selective also of cultural significance—they
is an uncommon subspecies advantage to the bears when they are considered a “spirit” animal
of the widespread black bear are stalking that prey. In terms by the local First Nations, whose
(U. americanus); about one-tenth of nutrient cycling, the migrating mythology includes stories about
of the black bear population has salmon are a conveyor of marine them, and they are also the official
a whitish pelage. Like other black nitrogen and phosphorus to the animal of British Columbia.
bears, animals in this population
are omnivorous in the feeding,
Copyright 2015 Nelson Education Ltd. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. Due to electronic rights, some third party content may be suppressed from the eBook and/or eChapter(s).
Nelson Education reserves the right to remove additional content at any time if subsequent rights restrictions require it.
To the natural world—may it ever be sustained and
complete.
–BF AND DG
To my dear little girl, Alexandra.

–JH
To my students, who have inspired me far more than

I could ever have inspired them.
–JPS
To Petra.
–RS
To my dear wife, Evelyn, who encouraged me to participate

in this project and then willingly relinquished endless
hours of together-time to allow me to complete it.
–RT
To Drexel, who taught me about perseverance in the

face of difficulty.
–RLW
To Jack Dainty and Bob Jefferies, who taught me

plant physiology.
–DRB
BRIEF CONTENTS
About the Authors xi
Your Tour of Ecology xxii
CHAPTER 1 Introduction to Ecology 2
CHAPTER 2 Environmental Influences 28
CHAPTER 3 Ecological Energetics 44
CHAPTER 4 Nutrients and Their Cycling 70
CHAPTER 5 Population Ecology 94
CHAPTER 6 Behavioural Ecology 134
CHAPTER 7 Physiological Ecology 158
CHAPTER 8 Life Histories 212
CHAPTER 9 Community Ecology 248
CHAPTER 10 Disturbance and Succession 296
CHAPTER 11 Biomes and Ecozones 338
CHAPTER 12 Biodiversity 368
CHAPTER 13 Landscape Ecology 392
CHAPTER 14 Conservation of the Natural World 430
CHAPTER 15 Resource Ecology 478
CHAPTER 16 Paleoecology: Lessons from the Past 534
CHAPTER 17 Ecology and Society 566
References 597
Glossary 621
Name Index 639
Species Index 643
Subject Index 647
NEL BRIEF CONTENTS v

TABLE OF CONTENTS
About the Authors xi CHAPTER 3
Your Tour of Ecology xxii
Ecological Energetics
3.1 Energy in Ecosystems 45
CHAPTER 1
3.2 Fundamentals of Energy 46
Introduction to Ecology States of Energy 46
3.3 Laws of Thermodynamics 49
1.1 Foundations of Ecology 3
First Law of Thermodynamics 49
Ecology and the Natural World 3
Second Law of Thermodynamics 49
Organisms and Environment 5
Fundamentals 5 3.4 Energy Flows and Budgets 50
Ecology within the Hierarchy of Organization 7 3.5 The Greenhouse Effect 53
Connections and Constraints 8
Ecology in Depth 3.1: Warming the Tundra 56
The Biosphere 10
1.2 The Science of Ecology 13 3.6 Energy Fixation in Ecosystems 57
Autotrophs 57
Ecological Data 15
A Canadian Ecologist 3.1: Suzanne Dufour: Chemosymbiosis—A
Subject Areas of Ecology 15
Win-Win Relationship 58
Ecologists 17
A Canadian Ecologist 1.1: Chris Pielou: Mathematical Ecology and Heterotrophs 59
More 18 Ecological Productivity 60
Ecology in Depth 3.2: Conserving the Carbon Stocks of Natural
The Scientific Method 18
Ecosystems 63
Environmental Applications 1.1: Environmental Gradients Near a
Metal Smelter 22 3.7 Food Webs 64
1.3 Evolutionary Ecology 23 Inefficiency of Energy Transfer 66
A Canadian Ecologist 3.2: Mike Apps: Carbon in Ecosystems 68
1.4 Ecology and Sustainability 25 CHAPTER SUMMARY 69
CHAPTER SUMMARY 27 QUESTIONS FOR REVIEW AND DISCUSSION 69
QUESTIONS FOR REVIEW AND DISCUSSION 27
CHAPTER 4
CHAPTER 2
Nutrients and Their Cycling
Environmental Inf luences 4.1 Nutrients 71
2.1 Environmental Influences 29 Ecology in Depth. 4.1: Toxicity and Evolution 72
Biotic and Abiotic Influences 29 4.2 Nutrient Cycles 74
Environmental Influences May be Optimal
Chemical and Biological Transformations 76
or Suboptimal 30
The Principle of Limiting Factors 31 4.3 The Carbon Cycle 77
Ecology in Depth 2.1: Extreme Environments 32 A Canadian Ecologist 4.1: Catherine Potvin: Studies of Global
Environmental Applications 2.1: Limiting Factors and the Cause of Change 80
Eutrophication 33 4.4 The Nitrogen Cycle 80
2.2 Environmental Stressors 35 Nitrogen Fixation 81
Ammonification and Nitrification 82
Kinds of Environmental Stressors 36
Denitrification 83
Environmental Applications 2.2: Chemicals That Are Potentially
Toxic: Contamination, Pollution, and Biomagnification 37 4.5 The Phosphorus Cycle 83
Environmental Applications 2.3: Chemical Stressors at Extremely 4.6 The Sulphur Cycle 84
Low Exposures: Endocrine Disruptors in Ecosystems 38 Environmental Applications 4.1: Pollution by Nutrients 85
Tolerance, Resilience, and Stability 38 4.7 Base Cations 86
Ecology in Depth 2.2: Natural Pollution at the
Smoking Hills 39 4.8 Soil as an Ecosystem 87
Physical Properties of Soil 88
2.3 Ecological Responses to Changes in Environmental Soil Types 90
Stress 40 Soil Degradation 90
A Canadian Ecologist 2.1: Tom Hutchinson: An Environmental A Canadian Ecologist 4.2: David Patriquin: A Champion of Organic
Ecologist 42 Management 92
CHAPTER SUMMARY 42 CHAPTER SUMMARY 93
QUESTIONS FOR REVIEW AND DISCUSSION 43 QUESTIONS FOR REVIEW AND DISCUSSION 93
NEL
vi
CHAPTER 5 Ecology in Depth 6.2: Conflict between Selection on Genes and on
Individuals 137
Population Ecology 6.2 Foraging for Food 137
5.1 Models in Ecology 95 Foraging While Threatened by Enemies 138
6.3 Natural Enemies and Adaptive Behaviour 138
5.2 Population Change 96
6.4 Interacting with Sexual Competitors 140
5.3 Exponential Population Growth 97
Why Is Sexual Competition Greater in Males? 141
Density-Independent per Capita Growth Rate (rmax) 97 Direct Competition between Rivals 142
Continuous-Time, Exponential Population Growth 98 Mating Preferences 144
Ecology in Depth 5.1: Linear versus Exponential Growth 99 The Sexes in Conflict 145
Discrete-Time, Exponential Population Growth 99 A Canadian Ecologist 6.1: Bridget Stutchbury: Behaviour, Ecology,
Relevance of r to Evolutionary Ecology, Resource Management, and Conservation of Migratory Songbirds 146
and Conservation 100 Sexual Selection Continues after Copulation 146
5.4 Logistic Population Growth 100 6.5 Social Behaviour 147
Density-Dependent per Capita Growth Rate (r) 101 Mutually Beneficial Acts and the Advantages of
Time Lags and Oscillations of Abundance 102 Group Life 147
Population Growth Rate and Sustainable Rates Altruism: Why Do Some Organisms Help Others? 148
of Harvesting 104 Ecology in Depth 6.3: Methodology in Behavioural Ecology: The
5.5 Age- and Stage-Structured Population Social Behaviour of Kin Selection 150
Growth 106 6.6 Social Selection 154
Age-Specific Schedules of Fecundity and Survival 106
Life Tables 107 6.7 Plant Behaviour 154
Realized per Capita Growth Rate on a Generational 6.8 Applications of Behavioural Ecology:
Time Scale 109 Community Ecology, Conservation, and
Realized per Capita Growth Rate in Continuous Time 109
Ecology in Depth 5.2: Derivation of the Euler-Lotka Equation 110
Human Behaviour 155
Predicting Changes in Abundance in an Age-Structured 6.9 Future Directions 156
Population 111 CHAPTER SUMMARY 156
Predicting Changes in Abundance in a Stage-Structured
Population 113
Application: The Collapse of Northern Atlantic Cod 114
Reproductive Value 115
5.6 Interspecific Competition 116
CHAPTER 7
The Lotka-Volterra Competition Model 116 Physiological Ecology
Competition in the Laboratory 117
Equilibrium Conditions: Isoclines and State-Space Graphs 118 7.1 Introduction to Physiological Ecology 159
Competitive Exclusion and Stable Coexistence: Graphical
Solutions 119
7.2 Physiological Ecology of Animals 160
Competitive Exclusion and Stable Coexistence: Algebraic Thermobiology 160
Solutions 120 Ecology in Depth 7.1: Scaling and Metabolism 162
Character Displacement 122 Ecology in Depth 7.2: Torpor in Bats 170
5.7 Predator–Prey Interactions 123 Ecology in Depth 7.3: The Convergent Evolution of Cryoprotectants
Influence of Predator–Prey Interactions on Population in Fish: Similar Solutions to Similar Problems 171
Growth 124 Environmental Applications 7.1: The Mountain Pine Beetle and
Functional Responses of Predators 125 Antifreeze 172
Population Dynamics of Predator–Prey Interactions 127
Water and Ion Balance 175
Variations in Predator–Prey Isoclines 130
Ecology in Depth 7.4: Water Balance in the Kangaroo Rat 181
Altered Predator–Prey Interactions Resulting from
Overfishing 131 7.3 Gas Exchange 182
A Canadian Ecologist 5.1: Charles Krebs: An Ecologist’s Population Role of the Circulatory System 182
Ecologist 132 Acid–Base Balance 188
CHAPTER SUMMARY 132 7.4 Physiological Ecology of Plants 191
QUESTIONS FOR REVIEW AND DISCUSSION 133 Photosynthesis 191
Water Balance 194
A Canadian Ecologist 7.1: Diane Srivastava: Scaling up from
CHAPTER 6 Miniature Food Webs to Global Biodiversity 195
Heat and Cold 202
Behavioural Ecology 7.5 Phytohormones 205
6.1 Behavioural Ecology 135 Secondary Compounds 206
Ecology in Depth 7.5: Trade-Offs and Induced Chemical
Ecology in Depth 6.1: Research Questions and Levels of Analysis in
Defences 210
Behavioural Ecology 136
CHAPTER SUMMARY 210
Adaptation, Selection, and Behaviour 136
NEL TA B L E O F C O N T E N T S vii
CHAPTER 8 9.3 Conceptual Models of Community Structure 286
Top–Down versus Bottom–Up Regulation and Trophic
Life Histories Cascades 286
Connell’s Model 289
8.1 Fundamentals of Life History Theory 213 Menge and Sutherland’s Model 290
Life-History Traits 215 Ecology in Depth 9.2: The Kluane Boreal Forest Ecosystem
Trait Variability 215 Project 291
Linking Life-History Traits to Population Growth CHAPTER SUMMARY 292
Rate 217 QUESTIONS FOR REVIEW AND DISCUSSION 293
Bet Hedging 220
8.2 Trade-Offs and the Costs of Reproduction 222
Trade-Offs 222 CHAPTER 10
Reproductive Constraints Impose Reproductive Costs 224
8.3 Natural Selection on Life Histories 226 Disturbance and Succession
Age at Maturity and Reproductive Effort 227 10.1 Disturbance and Succession 297
Influence of Growth Rate on Life History 230
r- and K-Selection 230 Resistance and Resilience 297
A Plant-Focused Classification 232 10.2 Characteristics of Disturbance 298
Ecology in Depth 8.1: The Fallacy of Fecundity and Extinction The Scale of Disturbance 299
Risk in Marine Fish 233 The Frequency of Disturbance 302
Life-History Invariants 234 Anthropogenic Disturbances 303
8.4 Offspring Size and Number 234 Environmental Applications 10.1: Alien Invaders as Biotic
Disturbances 304
Optimal Egg Size 235
8.5 Alternative Life Histories 238 10.3 Patterns and Mechanisms of Succession 305
Environmentally Determined Tactics and “The Best of a Late-Stage Communities 306
Bad Situation” 238 Alternative Stable States 309
Ecology in Depth 8.2: Extreme Alternative Life Histories 239 Mechanisms of Succession 311
Life History and Succession 312
Genetically Determined Strategies and Frequency-Dependent Means of Regeneration 312
Selection 240 Changes in Structural and Functional Properties
Threshold Traits: An Interaction between Environment and during Succession 314
Genotypes 240
10.4 Primary Succession 317
8.6 Harvest-Induced Evolution of Life History 241
Deglaciation 317
Changes in Age and Size at Maturity Caused Sand Dunes 321
by Harvesting 241 Ice-Scouring 323
A Canadian Ecologist 8.1: Derek Roff: Life Histories and
Genetics—From Fish to Crickets 242
10.5 Secondary Succession 324
Wildfire 324
Consequences of Harvest-Induced Changes in
Irruptive Animals and Diseases 327
Life History 243
A Canadian Ecologist 10.1: Yves Bergeron: Succession in the Eastern
CHAPTER SUMMARY 245
Boreal Forest 328
Timber Harvesting and Management 330
Environmental Applications 10.2: Emulation Silviculture: What Can
Forestry Learn from Wildfire? 332
CHAPTER 9 Abandoned Land 334
CHAPTER SUMMARY 336
Community Ecology
9.1 The Nature of the Community 249
Community Organization 250
Centrifugal Organization 254 CHAPTER 11
Community Composition 254
Ecology in Depth 9.1: The Importance of Fundamental Biomes and Ecozones
Research 258
11.1 Ecosystems at a Global Scale 339
9.2 Community Structure 258 Order Out of Complexity 339
Patterns and Processes 259 Biomes 340
The Niche 260 11.2 The Major Biomes 343
Interactions among Species 263
Terrestrial Biomes 344
Effects of Competition on Community Structure 265
Ecology in Depth 11.1: Old-Growth Ecosystems 346
Effects of Facilitation on Community Structure 269
Effects of Herbivory, Predation, and Disturbance on Freshwater Ecosystems 350
Community Structure 270 Marine Biomes 352
A Canadian Ecologist 9.1: Tony Sinclair: A Champion Anthropogenic Ecosystems 355
of Biodiversity 281 11.3 Ecozones and Ecological Regions 357
Effects of Chance, Rare, and Uncontrollable Events 281 Terrestrial Ecozones of Canada 360
Interactive Effects 283 Marine Ecozones of Canada 365
NEL
viii TA B L E O F C O N T E N T S
A Canadian Ecologist 11.1: Stan Rowe: A Specialist in Biomes and a 13.7 Tools of Landscape Ecology 420
Scientist-Citizen 366
Remote Sensing 420
CHAPTER SUMMARY 367 Geographical Information Systems 421
QUESTIONS FOR REVIEW AND DISCUSSION 367 Environmental Applications 13.2: Putting It All Together: Landscape
Ecology Helps Transdisciplinary Research 422
Modelling 423
CHAPTER 12 13.8 Who Uses Landscape Ecology? 423
Achievements, Problems, and the Way Forward 423
Biodiversity
Environmental Applications 13.3: Landscape-Scale Planning by the
12.1 Biodiversity 369 Nature Conservancy of Canada 424
Genetic Variation 370 A Canadian Ecologist 13.2: Marie-Josée Fortin: A Landscape
Ecology in Depth 12.1: Endemic Species 372 Ecologist 426
Richness of Species 374 CHAPTER SUMMARY 427
Ecology in Depth 12.2: A Census of Marine Life 376 QUESTIONS FOR REVIEW AND DISCUSSION 428
Richness of Communities 379
12.2 The Importance of Biodiversity 380
Instrumental Value 380 CHAPTER 14
Provision of Ecological Services 381
Aesthetic Value 383 Conservation of the Natural World
Intrinsic Value 383
Ecology in Depth 12.3: Influences of Biodiversity on Ecosystem
14.1 Conservation of the Natural World 431
Functions 384 The Meaning of Conservation 431
The Natural World 432
Biodiversity Is Viewed as Important 384
12.3 Measuring Biodiversity 385 14.2 The Biodiversity Crisis 433
Natural Extinctions 433
A Canadian Ecologist 12.1: John Macoun and Paul Hebert: Two
Anthropogenic Extinctions and Endangerment 435
Cataloguers of Canadian Biodiversity—The Old and the New 386
Environmental Applications 14.1: Feral Cats: Natural
Genetic-Level Biodiversity 387 Born Killers 448
Diversity of Species 387
Richness of Communities 389 14.3 Canadian Biodiversity at Risk 449
Ecology in Depth 12.4: Calculating Species Diversity 390 Canadian Species at Risk 449
CHAPTER SUMMARY 391 Communities at Risk 451
QUESTIONS FOR REVIEW AND DISCUSSION 391 14.4 Conservation Biology 452
Fields within Conservation Biology 452
Important Concepts in Conservation Biology 453
CHAPTER 13 Conservation Priorities 455
14.5 Protected Areas 457
Landscape Ecology Planning for Conservation 457
Design of Protected Areas 459
13.1 Introduction to Landscape Ecology 393
Environmental Applications 14.2: The Nature Conservancy of
The Importance of Landscape Ecology 395 Canada: Conservation Science in the Private Sector 460
Origins of Landscape Ecology 396
Kinds of Protected Areas 461
13.2 Structure of Landscapes 397 Extent of Protected Areas 462
Patches 397 Stewardship of Protected Areas 463
Environmental Applications 13.1: Using Geographical Information Environmental Applications 14.3: Burn, Baby, Burn: The Use of
Systems (GIS) 398 Prescribed Burns in Conservation Management 464
Corridors 401 14.6 Roles of Governments and Other Organizations
Networks 403
in Conservation 469
Mosaic and Matrix 403
13.3 Measuring and Characterizing Landscapes 406 14.7 Some Good News 470
Conservation Successes 470
13.4 Ecological Functions at Landscape Scale 407
A Canadian Ecologist 14.1: Mowat, Pimlott, Theberge, and
The Distribution of Water as a Landscape Function 407 Theberge: Dancing with Wolves 472
A Canadian Ecologist 13.1: Lenore Fahrig: Landscapes for
Ongoing Challenges 475
Sustainable Populations 408
CHAPTER SUMMARY 475
Disturbance as a Landscape Function 409 QUESTIONS FOR REVIEW AND DISCUSSION 476
13.5 Landscape Change 411
Understanding Temporal Change: The Historical Landscape 411
Understanding Temporal Change: The Landscape
Demographics Approach 414
CHAPTER 15
Understanding Change: Spatial Approaches 416
Resource Ecology
13.6 Scale in Space and Time 418
Scale in Space 418 15.1 Natural Resources and Ecological Sustainability 479
Scale in Time 418 Environmental Applications 15.1: Easter Island 481
Integration of Scale in Time and Space 419
Economics and Ecology 482
NEL TA B L E O F C O N T E N T S ix
Growth, Development, and Sustainability 485 Reconstructing Long-Term Trends in Pacific Salmon 557
Capital and Natural Resources 489 Historical and Modern Climate Change 559
Sustainability of the Human Enterprise 491 Ecology in Depth 16.4: Traditional Ecological Knowledge (TEK)
15.2 Harvesting and Managing Biological Meets Paleoecology: The Impacts of a Major Storm Surge on
Coastal Ecosystems of the Mackenzie River Delta 563
Resources 492
CHAPTER SUMMARY 565
Maximum Sustainable Yield 492
Regeneration 495 QUESTIONS FOR REVIEW AND DISCUSSION 565
Limitations on Productivity 496
Natural Mortality 497
Harvest-Related Mortality 498 CHAPTER 17
Unsustainable Harvesting 499
Why Are Potentially Renewable Resources Ecology and Society
Overharvested? 502
15.3 Resource Ecology in Practice 504 17.1 Ecology and Sustainability 567
Forests and Forestry 504 Fundamental Research 567
Marine Bioresources 516 Applied Research 568
A Canadian Ecologist 15.1: Myers, Hutchings, Pauly, Worm, and Ecology and Sustainability 568
Lotze: Speaking for the Fish 526 A Canadian Ecologist 17.1: David Schindler: A Champion of Ecology
and Environment 570
15.4 Integrated Resource Management 530
17.2 Ecological Integrity 572
CHAPTER SUMMARY 531
QUESTIONS FOR REVIEW AND DISCUSSION 532 17.3 Monitoring and Research 576
Environmental Indicators 576
The Relationship between Research and Monitoring 580
CHAPTER 16 17.4 Ecological Change and Damage 583
A Canadian Ecologist 17.2: Anne Salomon: Ecology and Sustainable
Paleoecology: Lessons from the Past Solutions 586
16.1 What is Paleoecology? 535 17.5 Environmental Impact Assessment 587
Ecology in Depth 16.1: Paleoecology of Dinosaurs by Philip J. Currie Planning Options 588
(University of Alberta) 536 Environmental Applications 17.1: Cumulative
The Importance of Paleoecology 536 Environmental Effects 589
A Canadian Ecologist 16.1: Philip J. Currie: Dinosaur Examples of Impact Assessments 590
Sleuth 538 17.6 Ecology as a Career 592
16.2 Environmental Change 539 17.7 Conclusions 594
Ecology in Depth 16.2: The Fossil Forest of Axel CHAPTER SUMMARY 594
Heiberg Island 540 QUESTIONS FOR REVIEW AND DISCUSSION 595
16.3 The Toolkit 541
Tree Rings 542
Sediment: A Window on the Past 544 References 597
Ecology in Depth 16.3: The Poop on Chimney Swifts 550 Glossary 621
16.4 Paleoecology and Environmental Issues 553 Name Index 639
Acidification of Lakes 554
Species Index 643
Aquatic Osteoporosis: Calcium Declines in Lakes 555
Eutrophication of Lakes: The Problem of Overfertilization 556 Subject Index 647
NEL
x TA B L E O F C O N T E N T S
ABOUT THE AUTHORS
Bill Freedman is an ecol- University Biology Chairs. Beyond academics, Bill is a

ogist and environmental squash player, a collector of folk art and natural artifacts,
scientist. He was born in and an enthusiastic naturalist and traveller who loves to
downtown Toronto and did spend time in wild places.
all of his schooling there.
He received his M.Sc. and Jeffrey A. Hutchings was
Ph.D. degrees from the born in Orillia, Ontario. He
University of Toronto, where made his initial environ-
he studied in the Department mental forays into the land-
Sheldon Bowles
of Botany. He has taught in scapes of nearby Shield

the Department of Biology at lakes and later the out-
Dalhousie University, Halifax, ports of Newfoundland.
Nova Scotia, since 1979. His maturing interests in
Jeff Hutchings
Bill Freedman
The conceptual frame- ecology ultimately stemmed
work of Bill’s research has been the influence of from field experiences ran-
environmental stressors on biodiversity and other ging from mountainous ter-
structural and functional attributes of ecosystems. Jeffrey A. Hutchings rain near Kispiox, British
Understanding the influence of stressor regimes is of Columbia, to lakes and rivers
theoretical interest, and it also helps guide the adjoining Georgian Bay, Ontario, and from interior and
management of ecological damage caused by disturbance coastal Newfoundland and Labrador to high-latitude lakes
and pollution. Bill’s research has examined the effects of on Baffin and Ellesmere Islands in Nunavut. His current
a wide range of industrial activities, with somewhat of a research centres on questions related to life history evolu-
tion, behavioural ecology, phenotypic plasticity, population
focus on forestry practices, as well as work on the
dynamics, and conservation biology of marine and anad-
environmental effects of acidification, eutrophication,
romous fishes, particularly Atlantic cod (Gadus morhua),
metals, pesticides, and sulphur dioxide. Other interests
Atlantic salmon (Salmo salar), and brook trout (Salvelinus
include carbon storage in ecosystems, urban ecology,
fontinalis). From an applied perspective, this work has
Arctic ecology, the biodiversity of Sable Island, the design bearing on questions pertaining to the depletion, recovery,
of environmental monitoring programs, and ecologically and sustainable exploitation of marine fish; interactions
sustainable systems of resource harvesting. More than between wild and farmed Atlantic salmon; the biodiversity
100 refereed publications in scientific journals have of Canadian fish; and the communication of science to
resulted from this work, plus several hundred book and decision makers and society.
encyclopedia chapters, research reports, and other Jeff is appointed as a Faculty of Science Killam
documents. Professor of Biology at Dalhousie University and
Bill is also engaged in developing curriculum Professor II in the Centre for Ecological and Evolutionary
materials in support of higher education, and as part of Synthesis at University of Oslo, Norway. He received his
that work he has written several textbooks, including B.Sc. from the University of Toronto and his M.Sc. and
Environmental Ecology (2nd edition, 1995), and Ph.D. from Memorial University of Newfoundland, and
Environmental Science: A Canadian Perspective (5th undertook postdoctoral research at the University of
edition, 2010). Bill has served on the board of directors of Edinburgh and Fisheries and Oceans Canada in
the Nature Conservancy of Canada since 1992, was the St. John’s, Newfoundland. In addition to Jeff’s teaching
chair of that board from 2007 to 2009, and wrote a and research responsibilities, he has served on many
retrospective of the organization (A History of the Nature advisory and administrative committees for non-
Conservancy of Canada, 2013). He has also participated in governmental and governmental organizations. He was
environmental impact assessments of proposed and a member (2001–2012) and chair (2006–2010) of the
operating industrial facilities, and has served on advisory Committee on the Status of Endangered Wildlife in
panels to government. In 2006, Bill received a Canadian Canada (COSEWIC), the national independent body
Environment Award, Gold Medal Level, in the category of responsible for advising the federal government on the
Community Awards for Conservation, from the Canadian status of species at risk in Canada. From 2009 to 2012, he
Geographic Society. In 2007, he received a Careeer chaired the Royal Society of Canada Expert Panel on
Achievement Award from the Canadian Council of Ocean Health and Marine Biodiversity. In 2012 and 2013
NEL ABOUT THE AUTHORS xi

he served as the fourth president of the Canadian Society Environmental Change. He received a B.Sc. from McGill
for Ecology and Evolution. He has been a member of the University, an M.Sc. from Brock University, and a Ph.D.
board of directors of WWF Canada since 2012. from Queen’s University. Following postdoctoral work in
the High Arctic with the Geological Survey of Canada, he
Darryl T. Gwynne was became a faculty member at Queen’s University. He has
born in Bristol, also held adjunct appointments at several other universi-
England, and moved to ties in Canada, the United States, and China.
Canada in 1966. He is John founded the Paleoecological Environmental
a professor of biology Assessment and Research Lab (PEARL) in 1991—a group
at the University of of about 30 researchers dedicated to the study of global
Darryl Gwynne
Toronto in Mississauga. environmental change, focusing primarily on lake

He received a B.Sc. in ecology. An ISI Highly Cited Researcher, he has authored
Biology at the University about 470 journal publications and book chapters, and
Darryl T. Gwynne
of Toronto in 1974 completed 19 books, including his textbook Pollution of
and a Ph.D. in Zoology and Entomology from Colorado Lakes and Rivers: A Paleoenvironmental Perspective, now in
State University in 1979. After conducting postdoctoral
its second edition and also being translated into Chinese.
research at the University of New Mexico, in 1981 he
He has lectured on all seven continents, including as the
took up a Queen Elizabeth II Research Fellowship at the
2008 Rutherford Lecturer at the Royal Society (London).
University of Western Australia. Since 1987 he has been
He was the founding editor of the international Journal
at the University of Toronto, where he currently teaches
animal behaviour and a fourth year “Topics in Ecology and of Paleolimnology (1987–2007), is the editor of the journal
Evolution” course. Environmental Reviews, is editor of the Developments in
Darryl’s research seeks to understand the factors that Paleoenvironmental Research book series, and is on the
control sexual selection and the “typical” sexual editorial boards of additional journals.
differences in behaviour and structure, such as ornaments Since 1990, he has received over 45 national and
and weapons. His studies and those of his students have international research and teaching awards, including an
investigated the consequences of mating systems as NSERC Steacie Fellowship, the 1992 Steacie Prize
diverse as those with extreme sexual selection on males (awarded to Canada’s top young scientist or engineer), a
(harem defence and male weaponry in New Zealand weta Canada Council Killam Fellowship, the Geological
(a group of cricket relatives) to the key study species, Association of Canada Past-Presidents’ Medal, the
those that are rare examples of female competition for Botanical Society of America Darbaker Prize, the NSERC
mates. The common element in the life histories of most Herzberg Gold Medal as Canada’s top scientist or
of the study species are important goods and services engineer (2004), the Rigler Prize from the Society of
offered by the males, such as prey used by dance flies, and Canadian Limnologists, the Royal Society of Canada
specialized glandular secretions in other insects. The Miroslaw Romanowski Medal for advances in the
value of these gifts has led—for some species—to role environmental sciences, an NSERC Award of Excellence,
reversals in behaviour, with females competing for access and the American Society of Limnology and Oceanography
to gift-bearing males and occasionally the evolution of Hutchinson Award. The Royal Canadian Geographical
male-like ornamentation in females. Society named him as the 2008 Environmental Scientist
Darryl is author of over 100 scientific papers, several of the Year (an honour shared with his brother, Jules
popular articles, and a book (Katydids and Bush-Crickets: Blais, of the University of Ottawa). In 2009, he was
Reproductive Behavior and Evolution of the Tettigoniidae, presented with the Killam Prize for the Natural Sciences
Cornell University Press, 2001). He is several chapters into from the Canada Council, as well as the Premier’s
a new book, a popular tome on Discovery Award for Life Sciences and Medicine. In 2013,
sexual selection that focuses he was awarded the inaugural Science Ambassador
on the life histories of insects Award and the 2013 Weston Family Prize for Lifetime
found in the area of his home Achievement in Northern Research. He has received
in the Credit River Valley. three honorary doctorates, from St. Francis Xavier
University, the University of Helsinki, and the University
John P. Smol is a professor of Waterloo.
in the Department of Biology John has received 10 teaching and scientific outreach
at Queen’s University, with awards from Queens University as well as outside
Bernard Clark
a cross-appointment to the agencies, including a 3M National Teaching Fellowship

School of Environmental in 2009, considered to be Canada’s top teaching honour.
Studies, where he also holds In 2010, following a nationwide search, Nature magazine
John P. Smol the Canada Research Chair in named John Canada’s Top Mid-Career Science Mentor.
NEL
xii ABOUT THE AUTHORS
In 2013, the Governor General named John an Officer of he did postdoctoral research at the University of Western
the Order of Canada. Ontario. Roy is a professor in the Department of Botany and
the Beaty Biodiversity Research Centre at the University
Roger Suffling came to of British Columbia in Vancouver. He teaches two under-
the School of Planning at graduate courses in ecology and an undergraduate and
the University of Waterloo graduate course in plant ecology. He is primarily an experi-
following the obtaining of mental field ecologist investigating population-level pro-
degrees in botany, ecology, cesses, such as competition and herbivory as influences
and weed science, and on community structure, specifically species diversity and
several years working in ecosystem function.
environmental consulting. Roy’s research, and that of his students, is supported
He teaches ecology, park primarily by the Natural Sciences and Engineering
Roger Suffling
planning, ecological policy Research Council (NSERC) and has been conducted in
making, environmental a wide range of community types in the boreal forest in
impact assessment, land- northern Canada, grasslands in western Canada and the
Roger Suffling scape ecology, and ecological United Kingdom, Garry oak ecosystems, the Negev
restoration. Roger’s research desert in Israel, riverine forests in the Serengeti
interests centre on the landscape ecology and manage-
National Park in Tanzania, and subtropical forests in
ment of boreal forests, including the role of forest fires
southern China.
and global warming effects on the ecology and economy
Roy has published more than 150 papers and book
of Canada’s mid-north. Equally, he and his students
chapters and has served on the editorial boards of Agro-
research the ecology and management of urban eco-
systems. Roger has published numerous papers on these Ecosystems, the Canadian Journal of Botany, and the Israeli
topics and has been a consultant for local, provincial, and Journal of Ecology and Evolution. He was on the editorial
national governments; a royal commission; private com- board of the Journal of Ecology for 20 years. One of his
panies; and First Nations. He chaired an Ontario govern- papers has been recognized by the British Ecological
ment scientific advisory committee on woodland caribou Society as one of the top 100 most influential papers ever
conservation, and for over 30 years has participated in published by the organization (1913–2012). In pursuit of
numerous environmental assessments and conservation his academic career, Roy and his wife have spent
issues in northern Ontario. Many of these activities focus sabbaticals in Wales, Turkey, Northern Ireland, Israel,
on applying ecological principles to management of China, and Argentina.
incremental landscape change, whether caused by urban In his spare time, Roy has a keen amateur interest in
growth, forest fires, recreation, or forest harvesting. Middle Eastern and Biblical archaeology. He and his wife
Roger has travelled to over 20 countries in his study of Evelyn are avid travellers and enjoy the outdoors. Together
ecology but has a special passion for the ecosystems and they have trekked the Annapurna circuit, the Mount
people of northern Ontario. For a decade, through the Everest base camp, the High Atlas, the Inca trail, and
Quetico Foundation, he co-organized a youth program
climbed Kilimanjaro. For many years they have been
for northern Ontario students to conduct landscape-scale
involved in the AWANA children’s ministry. Roy and
research in Quetico Provincial Park, a large wilderness
Evelyn have two married children, Alistair and Andrea,
reserve. Roger’s hobbies include a garden that manages
to manage itself, family history, cross-country skiing, and four grandchildren—and by the time this second
and wilderness canoeing. As an immigrant to Canada, edition is published, they will have five.
Roger is profoundly grateful for the gifts that this vast
and magnificent country has afforded him in friend- Richard L. Walker completed
ships and beautiful landscapes. He is concerned that his Ph.D. in Animal Physiology
the next generation of Canadians must build a more at Michigan State University
sustainable society. in 1975 and immediately acc-
epted a faculty position at
Roy Turkington was born the University of Calgary in
in Northern Ireland. After the Department of Biological
earning his B.Sc. (Hons) Sciences. Richard retired in
degree in Biological and June 2012 and is now enjoying
Richard Walker
life as an Instructor Emeritus.

Roy and Evelyn Turkington
Environmental Studies from

the New University of Ulster While in graduate school,
and his doctorate from the Richard developed a keen
Richard L. Walker interest in environmental phy-
University College of North
Wales, Roy immigrated with siology and the effects of pollutants on aquatic animals.
Roy and Evelyn Turkington his wife to Canada where He has had the privilege of working with some of Canada’s
NEL ABOUT THE AUTHORS xiii

leading environmental physiologists and has published Dawn trained as an ecologist, in the field of plant–
journal articles on aluminum toxicity in brook trout. herbivore interactions, and has carried out field research
Richard loves to teach and has received several in grasslands and forests, from temperate to arctic
teaching excellence awards from the Faculty of Science regions. Dawn has a B.Sc. in Biogeography and
and from the Students’ Union, and was recently inducted Environmental Studies and an M.Sc. in Botany from the
in Student’s Union Teaching Excellence Hall of Fame. He University of Toronto. Her D.Phil. in Zoology, from
is currently a sessional instructor in Human Physiology Oxford University’s Edward Grey Institute in Field
in the Faculty of Kinesiology. Ornithology, looked at sheep grazing behaviour.
As an avid hiker, backpacker, and skier, Richard Since joining York University, Dawn has won the
considers himself very fortunate to live in the foothills Faculty of Science and Engineering teaching award (2003)
of the Canadian Rockies. He also makes forays to the and the President’s University-Wide Teaching Award
West Coast where he enjoys sailing with family and (2013). As well as publishing dozens of journal articles,
friends. chapters, conference proceeding papers, and technical
reports, Dawn wrote a textbook, Ecology and Control of
Dawn Bazely is professor Introduced Plants: Evaluating and Responding to Invasive
of Biology in the Faculty of Plants, (2003, with Judith Myers), and coedited
Science at York University, Environmental and Human Security in the Arctic (2013).
Toronto, where she has After nearly 20 years away from Arctic fieldwork, Dawn
taught since 1990. She is returned to the north in 2002 for fieldwork in Sweden. In
also Director of IRIS, the
2006, she led the Canadian section of the International
university-wide Institute for
Polar Year project: GAPS (Gas, Arctic Peoples, and
Research and Innovation in
Security).
Sustainability (2006–11 and
2012–13). At IRIS, Dawn’s
In 2011 Dawn received a Charles Bullard Fellowship
from Harvard University, and spent her sabbatical at
Dawn Bazely
mission is to develop, lead,

and support interdisci- Harvard Forest, where she worked on a book examining
plinary research on diverse conservation and ecological issues in southern Ontario,
Dawn Bazely fronts. The student-led from scientific, policy, and political perspectives. This is
annual campus sustain- the most heavily populated, industrialized, and farmed
ability survey that she developed in 2007 is now a regular region of Canada, and it provides a case study for
event. These surveys of the York community explore evaluating and responding to what lies ahead for other
diverse topics ranging from climate change to local food parts of the temperate forest region in the face of climate
and transportation preferences. change and its associated issues and challenges.
NEL
xiv ABOUT THE AUTHORS
PREFACE
The Context of Ecology living in or at the edge of wilderness. Today Canada supports
Ecology is a vital way of knowing. This is because its an astonishing cosmopolitan and multicultural society, but
knowledge and predictions help us to understand how we still appreciate the natural world, as witnessed by its
the natural world functions, the station of humans within prominence in our art, coinage, literature, stamps, video,
that domain, and the means by which our use of its and other cultural expressions. Many of our students are
resources can be undertaken on a sustainable basis. For newcomers to this country, and great numbers have been
these reasons, the wisdom of ecology can beneficially raised exclusively in urban environments. It is vitally
inform key aspects of our society and economy. important that they mature with a solid understanding of
Ecology has a global base of data and understanding the ecological life support system, whether of the vast tundra
and a universal set of principles that are relevant any- and forest of the North or the more familiar parks and back-
where on Earth. A goal of any introductory textbook in yards of Canada’s cities.
ecology is to facilitate an understanding of that core of its Notwithstanding the apparent high regard with
knowledge. However, ecology also involves the study of which the peoples of Canada have always viewed the nat-
wild species and their higher-level aggregations, such as ural world, many of our native species and natural eco-
populations, communities, and landscapes. In that sense, systems are at great risk of disappearing. This damage
ecology has a profound spatial context that extends from has been caused by extensive conversions of natural habi-
relatively local situations to much larger regional scales, tats into human-dominated land uses, and by degradation
and ultimately, to the entire biosphere. caused by alien species and diseases, excessive resource
Within that context, however, the boundaries of harvesting, pollution, and other anthropogenic stressors.
countries are not ecoregional in their layout, meaning These are exceedingly important environmental dam-
they were not designed according to natural ecological ages, and also socioeconomic ones because they pose a
precincts. Nevertheless, national borders do specify par- grave threat to the longer-term sustainability of the Can-
ticular expanses of land and sea. That spatial fact provides adian economy. In large part those damages are ecological
an important context for the ways that ecology is taught in character, as are their mitigations.
and learned in any country.
In this sense, Canada supports particular arrange- A Canadian Approach
ments and dynamics of ecosystems and species, which are Clearly, there is a Canadian context to learning about
affected by the human economy in specific ways. The nat- ecology. This obvious deduction is the reason that we have
ural ecosystems of our country range from temperate Caro- chosen to develop this textbook—Ecology: A Canadian Con-
linian forest in extreme southern Ontario, to true desert in text, Second Edition—which you are now beginning to read.
the southern Okanagan Valley of British Columbia, to It is our belief that this new book will be helpful to Canadian
High-Arctic tundra on the islands of Nunavut. On the students as they seek to learn about ecology, but in the con-
Pacific coast there are humid temperate rainforests, while text of our country. We also believe that this textbook will
in the centre of Canada there are expansive montane and assist instructors of ecology in colleges and universities in
boreal forests and prairie grasslands, and on the Atlantic Canada, in ways that books developed in other countries
coast a mixture of temperate and boreal forests. The marine cannot do as well. We sincerely hope that you will appre-
realms range from frigid and ice-covered waters north of ciate, and benefit from, our efforts to achieve these goals.
Ellesmere Island to boreal and temperate ecosystems in It is important to recognize that this textbook has
more southerly reaches of the marine estate of Canada. The emerged from a process that is different from the more
ecological communities that occur in these far-flung eco- common procedure of “Canadianizing” versions of text-
regions of Canada, and the species they sustain, comprise books that were originally developed for use by students
the essence of the biodiversity of our country. and instructors in a different country, usually the United
The peoples of Canada have always esteemed the nat- States. Instead of taking the relatively easy path of modi-
ural values of their lands. First Nations and Inuit venerated fying an existing textbook by inserting some Canadian
animals and plants that were exploited as food, medicine, content, we have created a totally new one, from the ground
and materials, as well as the habitats that they all shared. An up. This book was specifically designed and written for
attitude of respect and awe of the natural world was also held Canadian students and their instructors in Canada, by Can-
by many early European and other immigrants to Canada, adians well experienced in the ecology of our country.
although those feelings may have been tempered by trepida- In essence, the approach we took was to identify the
tion because of their often precarious circumstances of core subject areas of ecology, which are relevant in all
NEL P R E FA C E xv
countries, and develop clear explanations of the basic increased coverage in Chapter 13, Landscape
principles. Wherever it was suitable, however, we illus- Ecology, and a new Environmental Applications
trated those basics using data and case material relevant Box 14.1—Feral Cats: Natural Born Killers.
to Canada, and usually derived from studies by Canadian
• Freshwater Ecology: We have enhanced the coverage
ecologists. This is not to say that our book does not also
of freshwater ecosystems throughout the book,
contain abundant international material, because it cer-
including new examples in Chapter 4, Nutrients and
tainly does. The key point is that the essential approach
Their Cycling, and Chapter 9, Community Ecology.
of our treatment of introductory ecology is a balanced
integration of global and Canadian contexts. • New Boxes: We have added an impressive number
of new boxes that explain key concepts in greater
Ecology, Evolution, and Sustainability detail, provide illustrative case material of the appli-
This textbook is about the fundamentals of ecology within cation of ecological knowledge, or feature the contri-
a Canadian context. However, evolution and sustainability butions of Canadian ecologists so that our students
also provide essential perspectives for the study of ecology, can understand our large national influence on the
and we have endeavoured to integrate these themes into global endeavour of ecology. Among these new
the book wherever they are relevant. The vitally important boxes is Ecology in Depth 16.1—The Paleoecology
subjects of evolution and evolutionary relationships are of Dinosaurs, contributed by Phil Currie, celebrated
examined across the curriculum, whenever they are paleontologist at the University of Alberta.
helpful in understanding concepts and case material. • Chapter Summaries: We have also now included end-
We have also used this approach to explore the vital of-chapter summaries, which list the key learning out-
intersection of ecology and sustainability and to integrate comes that students are expected to have mastered.
those links whenever subjects such as resource depletion,
pollution, and the conservation of biodiversity are exam- We sincerely hope that you will appreciate our hard
ined. In addition, there are two chapters that explore work in improving this textbook—in particular, because
these imperative themes more comprehensively—one we had your needs foremost in our minds.
about resource ecology, and the other about the conserva-
tion of the natural world. Organization
The field of ecology is wide and interdisciplinary, so much
New to the Second Edition so that it cannot all be covered in an introductory-level
We have worked hard to improve this second edition of class or in a textbook of a sensible length. When this book
the textbook. We engaged in a wide solicitation of and its second edition were being planned, we consulted
instructors from universities and colleges across Canada widely with colleagues and anonymous reviewers, who
to gather suggestions for improvements, and then dili- advised us that they wanted to see a textbook of modest
gently followed up with many constructive improvements length (and price) that covered the key themes of ecology,
in all chapters of the book. These improvements include: but not necessarily the entirety of its subject matter. To
achieve that goal, we divided the field into 17 chapters,
• Tone, Language, and Consistency: We completed a which are interconnected where relevant, but are still
comprehensive editorial review to improve the con-
organized in ways that make them suitable for teaching
sistency of language and tone and diminish redun- and learning as independent units.
dancy across the chapters. We have organized Ecology: A Canadian Context,
• Enhanced Instructional Imagery: We have greatly Second Edition, into the following 17 chapters:
enhanced the visual appeal of the book, with over
200 new and improved photos and illustrations that Chapter 1: Introduction to Ecology
illustrate key concepts. Bill Freedman
• Currency: We have thoroughly updated the treat- This chapter establishes the foundations of ecology, and
ment of all time-sensitive information and issues examines its methodology within the context of the prin-
across all of the chapters. ciples and practices of scientific investigation. One section
examines evolution as an overarching theme of ecology,
• Plant Ecology: In addition to enhancing our coverage of and another explains how ecological considerations are
plant ecology throughout the book, we have introduced vital to framing the sustainability of the human economy.
a greatly expanded chapter on physiological ecology
that includes a major new section on plants, creating a Chapter 2: Environmental Influences
more balanced treatment of that subject area. Bill Freedman
• Urban Ecology: We have expanded the coverage of Here we examine the environmental factors that influence
urban ecology throughout the book, including species and ecosystems. While noting that ecosystems are
NEL
xvi P R E FA C E
always subjected to pervasive change, we examine con- interplay of biological variation and natural selection. The
siderations that might result in change being viewed as principal subject areas examined are the costs of
ecological or economic damage. reproduction, life histories and fitness, trade-offs between
the numbers of offspring and their sizes, alternative life
Chapter 3: Ecological Energetics histories, and influences of anthropogenic harvesting.
Bill Freedman
This chapter explains the forms in which energy may exist Chapter 9: Community Ecology
and the laws of thermodynamics that govern their trans- Roy Turkington
formations. The energy budget of Earth is described, Ecological communities are examined as groups of spe-
including the vital greenhouse effect that helps to maintain cies that live together and interact, directly or indirectly,
the planet within a temperature range appropriate for life through competition for scarce resources, herbivory,
and ecosystems. We then examine ecological energetics, predation, disease, and facilitation. Environmental and
beginning with the fixation of solar energy by primary pro- biological influences on the structure and dynamics of
ducers, moving through the passage of fixed energy along communities are examined, including adaptive responses.
food webs, and ending with the decomposition process that
oxidizes dead organic matter to return degraded energy Chapter 10: Disturbance and Succession
and simple inorganic molecules to the environment. Bill Freedman
The causes and consequences of disturbances are explained,
Chapter 4: Nutrients and Their Cycling including those resulting from natural and anthropogenic
Bill Freedman influences. This is followed by consideration of the mech-
Nutrients are explained as substances needed for the healthy anisms and patterns of successional recovery, including
physiology and growth of organisms. The cycling of carbon, case studies of both primary and secondary succession.
nitrogen, phosphorus, and sulphur, and the bases calcium,
Chapter 11: Biomes and Ecozones
magnesium, potassium, and sodium are examined. We also
Bill Freedman
investigate soil as an ecosystem, including influences on the
The major terrestrial and marine biomes of the world are
formation of dominant kinds of soils.
described, in both the terrestrial and marine realms, as are
Chapter 5: Population Ecology key habitats such as types of wetlands and anthropogenic
Jeffrey A. Hutchings ecosystems. The terrestrial and marine ecozones of both
This chapter describes the ways that populations may vary Canada and the rest of North America are examined.
over time, including exponential and logistical changes and
Chapter 12: Biodiversity
their explanatory models. The influences of age structure,
Bill Freedman
competition, and trophic interactions on population change
The hierarchical levels of biodiversity are explained, begin-
are explored, including their evolutionary contexts.
ning with genetic variation, then species richness, and
Chapter 6: Behavioural Ecology extending to community-scale patches on landscapes and
seascapes. The ways of measuring biodiversity at these
Darryl T. Gwynne
scales are also examined. Biodiversity’s importance is
Here, we examine interactions among behaviour, ecological
explained, including its intrinsic value and the vital goods
relationships, and adaptive evolutionary change. The prin-
and services that are provided to humans and our economy.
cipal topics are foraging, defence against predators, sexual
selection, and the evolution of social behaviour. Chapter 13: Landscape Ecology
Roger Suffling
Chapter 7: Physiological Ecology
This chapter describes spatial approaches to the structure
Richard L. Walker and dynamics of landscapes, including both natural and
Dawn Bazely anthropogenic effects. The major ways of measuring the
This field is examined through the adaptive physiological attributes of landscapes are explored, including the use
traits of animals and plants in relation to their environmental of geographical information systems.
conditions. Particular attention is paid to thermobiology,
water and ionic balances, gas exchange and transport, acid– Chapter 14: Conservation of the Natural World
base balances, and influences of environmental stressors. Bill Freedman
The modern extinction crisis is examined and put into the
Chapter 8: Life Histories context of previous mass extinctions caused by natural
Jeffrey A. Hutchings forces. The basic tenets of conservation biology are
This chapter distills the diverse life histories of species explained, including the design and stewardship of pro-
into sets of responses that are results of the dynamic tected areas. Canada’s biodiversity at risk is described, as
NEL P R E FA C E xvii
are the conservation roles and responsibilities of govern- • Tables, Figures, and Photos. We have gone to great
ments and other organizations. The chapter ends with lengths to present easily digestible and abundant
success stories involving cases of endangered species that information in tables and figures, as well as plen-
have been rescued from the brink of extinction. tiful and attractive photos that illustrate concepts,
habitats, and species that are well communicated in
Chapter 15: Resource Ecology a visual medium.
Bill Freedman • Key Terms. Whenever we mention a word or phrase
Ecological economics are explained as a foundation for that is a core part of the lexicon of ecology, it is
understanding the concept of economic sustainability and highlighted and defined. All of these terms are
its fundamental reliance on renewable resources rather aggregated into a comprehensive glossary at the
than nonrenewable ones. Systems of harvesting and man- end of the book.
aging bioresources are explained, and international and
Canadian case material is presented to illustrate the phe- End-of-Chapter Learning and Review
nomenon of overharvesting. Improved management sys- Each chapter ends with a number of features that help to
tems that would allow sustainable use are described, cultivate learning of the subject matter. These are:
including integrated ones that accommodate both the eco-
nomic values of resources as well as ecological considera- • Chapter Summary. This is a bulleted list that sum-
tions such as biodiversity and environmental services. marizes the key concepts and learning outcomes
that students are expected to master in each
Chapter 16: Paleoecology: Lessons from the Past chapter. Each point is tied to one of the learning
John P. Smol objectives outlined at the beginning of the chapter.
Longer-term ecological changes are described, as are the • Questions for Review and Discussion. A number of
paleoecological methods that have allowed their causes questions are presented for students to answer.
and consequences to be examined. The importance of These are intended to assist in review of the subject
paleoecological studies to understanding environmental matter, while aiding comprehension and facili-
issues is also discussed. tating in-class discussion of certain topics. The
Instructor’s Manual for the book provides suggested
Chapter 17: Ecology and Society answers for all of these questions.
Bill Freedman
This final chapter examines the concept of ecological End-of-Book Features
integrity and the processes of ecological monitoring and The following resources are found at the end of the book:
research and environmental impact assessment. Eco-
logical sustainability is also explained, as is the import- • References. All references cited in the chapter are
ance of the knowledge and wisdom of ecology, and the listed in a comprehensive bibliography at the back
work of ecologists in guiding progress to that goal. of the book.
• Glossary. Definitions for all key terms are provided
Features of the Text in this essential resource. Page references are
In-Chapter Features and Learning Aids included to help students review these key terms in
The defining attributes of this textbook are the ways that the context of the original discussion.
it presents the fundamentals of ecology within a Canadian • Indexes. Three types of index are included for
context, while integrating the concepts of evolution and readers’ ease of reference. They consist of Name
sustainability wherever they are relevant. The book con- (of persons mentioned in the book), Species, and
tains several features that help to further these goals: Subject indexes.
• Boxed Features. All chapters have stand-alone
boxed elements that present detailed examinations Instructor Ancillaries
of selected important concepts or helpful case About the Nelson Education Teaching Advantage (NETA)
material. The three types of boxes are (1) Ecology The Nelson Education Teaching
in Depth, which is intended to provide a Advantage (NETA) program delivers
detailed investigation of important concepts; research-based instructor resources
(2) Environmental Applications, an in-depth look at that promote student engagement and higher-order
real-world applications of content discussed in the thinking to enable the success of Canadian students and
chapter; and (3) A Canadian Ecologist, which high- educators. To ensure the high quality of these materials,
lights prominent studies done by ecologists who all Nelson ancillaries have been professionally
have worked in Canada. copyedited.
NEL
xviii P R E FA C E
Be sure to visit Nelson Education’s Inspired Instruc- resulting focus on pedagogical utility, rather than
tion website at http://www.nelson.com/inspired to find technological wizardry, ensures that all of our tech-
out more about NETA. Don’t miss the testimonials of nology supports better outcomes for students.
instructors who have used NETA supplements and seen
student engagement increase! Instructor Resources
• Planning Your Course. NETA Engagement presents All NETA and other key instructor ancillaries are provided
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content and activities to their classes. NETA com/ecology2e, giving instructors the ultimate tool for
Instructor’s Manuals not only identify the topics customizing lectures and presentations.
that cause students the most difficulty, but also • NETA Test Bank. This resource includes multiple-
describe techniques and resources to help students choice questions written according to NETA guide-
master these concepts. Dr. Roger Fisher’s lines for effective construction and development of
Instructor’s Guide to Classroom Engagement accom- higher-order questions. The Test Bank was
panies every Instructor’s Manual. copyedited by a NETA-trained editor. Also included
• Assessing Your Students. NETA Assessment relates are true-and-false questions.
to testing materials. NETA Test Bank authors create • The NETA Test Bank is available
multiple-choice questions that reflect research- in a new, cloud-based platform.
based best practices for constructing effective ques- Testing Powered by Cognero®* is a secure online
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NEL P R E FA C E xix
Another random document with
no related content on Scribd:
“May eighth. This morning I reached the Cooper’s Creek depot
and found no sign of Mr. Burke’s having visited the creek, or the
natives having disturbed the stores.”
Only a few miles away the creek ran out into channels of dry
sand where Burke, Wills and King were starving, ragged beggars fed
by the charitable black fellows on fish and a seed called nardoo, of
which they made their bread. There were nice fat rats also, delicious
baked in their skins, and the natives brought them fire-wood for the
camp.
Again they attempted to reach the Mounted Police outpost, but
the camels died, the water failed, and they starved. Burke sent Wills
back to Cooper’s Creek. “No trace,” wrote Wills in his journal, “of any
one except the blacks having been here since we left.” Brahe and
Wright had left no stores at the camp ground.
Had they only been bushmen the tracks would have told Wills of
help within his reach, the fish hooks would have won them food in
plenty. It is curious, too, that Burke died after a meal of crow and
nardoo, there being neither sugar nor fat in these foods, without
which they can not sustain a man’s life. Then King left Burke’s body,
shot three crows and brought them to Wills, who was lying dead in
camp. Three months afterward a relief party found King living among
the natives “wasted to a shadow, and hardly to be distinguished as a
civilized being but by the remnants of the clothes upon him.”
“They should not have gone,” said one pioneer of these lost
explorers. “They weren’t bushmen.” Afterward a Mr. Collis and his
wife lived four years in plenty upon the game and fish at the
Innaminka water-hole where poor Burke died of hunger.
Such were the first crossings from east to west, and from south
to north of the Australian continent.
XVIII
A. D. 1867
THE HERO-STATESMAN
THERE is no greater man now living in the world than Diaz the
hero-statesman, father of Mexico. What other soldier has scored
fourteen sieges and fifty victorious battles? What other statesman,
having fought his way to the throne, has built a civilized nation out of
chaos?
This Spanish-red Indian half-breed began work at the age of
seven as errand boy in a shop. At fourteen he was earning his living
as a private tutor while he worked through college for the priesthood.
At seventeen he was a soldier in the local militia and saw his country
overthrown by the United States, which seized three-fourths of all
her territories. At the age of twenty-one, Professor Diaz, in the chair
of Roman law at Oaxaca, was working double tides as a lawyer’s
clerk.
In the Mexican “republic” it is a very serious offense to vote for
the Party-out-of-office, and the only way to support the opposition is
to get out with a rifle and fight. So when Professor Diaz voted at the
next general election he had to fly for his life. After several months of
hard fighting he emerged from his first revolution as mayor of a
village.
The villagers were naked Indians, and found their new mayor an
unexpected terror. He drilled them into soldiers, marched them to his
native city Oaxaca, captured the place by assault, drove out a local
usurper who was making things too hot for the citizens, and then
amid the wild rejoicings that followed, was promoted to a captaincy
in the national guards.
Captain Diaz explained to his national guards that they were fine
men, but needed a little tactical exercise. So he took them out for a
gentle course of maneuvers, to try their teeth on a rebellion which
happened to be camped conveniently in the neighborhood. When he
had finished exercising his men, there was no rebellion left, so he
marched them home. He had to come home because he was
dangerously wounded.
It must be explained that there were two big political parties, the
clericals, and the liberals—both pledged to steal everything in sight.
Diaz was scarcely healed of his wound, when a clerical excursion
came down to steal the city. He thrashed them sick, he chased them
until they dropped, and thrashed them again until they scattered in
helpless panic.
The liberal president rewarded Colonel Diaz with a post of such
eminent danger, that he had to fight for his life through two whole
years before he could get a vacation. Then Oaxaca, to procure him a
holiday, sent up the young soldier as member of parliament to the
capital.
Of course the clerical army objected strongly to the debates of a
liberal congress sitting in parliament at the capital. They came and
spoiled the session by laying siege to the City of Mexico. Then the
member for Oaxaca was deputed to arrange with these clericals.
He left his seat in the house, gathered his forces, and chased
that clerical army for two months. At last, dead weary, the clericals
had camped for supper, when Diaz romped in and thrashed them.
He got that supper.
So disgusted were the clerical leaders that they now invited
Napoleon III to send an army of invasion. Undismayed, the
unfortunate liberals fought a joint army of French and clericals,
checked them under the snows of Mount Orizaba, and so routed
them before the walls of Puebla that it was nine months before they
felt well enough to renew the attack. The day of that victory is
celebrated by the Mexicans as their great national festival.
In time, the French, forty thousand strong, not to mention their
clerical allies, returned to the assault of Puebla, and in front of the
city found Diaz commanding an outpost. The place was only a large
rest-house for pack-trains, and when the outer gate was carried, the
French charged in with a rush. One man remained to defend the
courtyard, Colonel Diaz, with a field-piece, firing shrapnel, mowing
away the French in swathes until his people rallied from their panic,
charged across the square, and recovered the lost gates.
The city held out for sixty days, but succumbed to famine, and
the French could not persuade such a man as Diaz to give them any
parole. They locked him up in a tower, and his dungeon had but a
little iron-barred window far up in the walls. Diaz got through those
bars, escaped, rallied a handful of Mexicans, armed them by
capturing a French convoy camp, raised the southern states of
Mexico, and for two years held his own against the armies of France.
President Juarez had been driven away into the northern desert,
a fugitive, the Emperor Maximilian reigned in the capital, and
Marshal Bazaine commanded the French forces that tried to conquer
Diaz in the south. The Mexican hero had three thousand men and a
chain of forts. Behind that chain of forts he was busy reorganizing
the government of the southern states, and among other details,
founding a school for girls in his native city.
Marshal Bazaine, the traitor, who afterward sold France to the
Germans, attempted to bribe Diaz, but, failing in that, brought nearly
fifty thousand men to attack three thousand. Slowly he drove the
unfortunate nationalists to Oaxaca and there Diaz made one of the
most glorious defenses in the annals of war. He melted the cathedral
bells for cannon-balls, he mounted a gun in the empty belfry, where
he and his starving followers fought their last great fight, until he
stood alone among the dead, firing charge after charge into the
siege lines.
Once more he was cast into prison, only to make such frantic
attempts at escape that in the end he succeeded in scaling an
impossible wall. He was an outlaw now, living by robbery, hunted like
a wolf, and yet on the second day after that escape, he commanded
a gang of bandits and captured a French garrison. He ambuscaded
an expedition sent against him, raised an army, and reconquered
Southern Mexico.
Porfirio Diaz
It was then (1867) that the United States compelled the French
to retire. President Juarez marched from the northern deserts,
gathering the people as he came, besieged Querétaro, captured and
shot the Emperor Maximilian. Diaz marched from the south, entered
the City of Mexico, handed over the capital to his triumphant
president, resigned his commission as commander-in-chief, and
retired in deep contentment to manufacture sugar in Oaxaca.
For nine years the hero made sugar. Over an area in the north
as large as France, the Apache Indians butchered every man,
woman and child with fiendish tortures. The whole distracted nation
cried in its agony for a leader, but every respectable man who tried
to help was promptly denounced by the government, stripped of his
possessions and driven into exile. At last General Diaz could bear it
no longer, made a few remarks and was prosecuted. He fled, and
there began a period of the wildest adventures conceivable, while
the government attempted to hunt him down. He raised an
insurrection in the north, but after a series of extraordinary victories,
found the southward march impossible. When next he entered the
republic of Mexico, he came disguised as a laborer by sea to the port
of Tampico.
At Vera Cruz he landed, and after a series of almost miraculous
escapes from capture, succeeded in walking to Oaxaca. There he
raised his last rebellion, and with four thousand followers
ambuscaded a government army, taking three thousand prisoners,
the guns and all the transport. President Lerdo heard the news, and
bolted with all the cash. General Diaz took the City of Mexico and
declared himself president of the republic.
Whether as bandit or king, Diaz has always been the
handsomest man in Mexico, the most courteous, the most charming,
and terrific as lightning when in action. The country suffered from a
very plague of politicians until one day he dropped in as a visitor,
quite unexpected, at Vera Cruz, selected the eleven leading
politicians without the slightest bias as to their views, put them up
against the city wall and shot them. Politics was abated.
The leading industry of the country was highway robbery, until
the president, exquisitely sympathetic, invited all the principal
robbers to consult with him as to details of government. He formed
them into a body of mounted police, which swept like a whirlwind
through the republic and put a sudden end to brigandage. Capital
punishment not being permitted by the humane government, the
robbers were all shot for “attempting to escape.”
Next in importance was the mining of silver, and the recent
decline in its value threatened to ruin Mexico. By the magic of his
finance, Diaz used that crushing reverse to lace the country with
railroads, equip the cities with electric lights and traction power far in
advance of any appliances we have in England, open great
seaports, and litter all the states of Mexico with prosperous factories.
Meanwhile he paid off the national debt, and made his coinage
sound.
He never managed himself to speak any other language than his
own majestic, slow Castilian, but he knew that English is to be the
tongue of mankind. Every child in Mexico had to go to school to learn
English.
And this greatest of modern sovereigns went about among his
people the simplest, most accessible of men. “They may kill me if
they want to,” he said once, “but they don’t want to. They rather like
me.” So one might see him taking his morning ride, wearing the
beautiful leather dress of the Mexican horsemen, or later in the day,
in a tweed suit going down to the office by tram car, or on his
holidays hunting the nine-foot cats which we call cougar, or of a
Sunday going to church with his wife and children. On duty he was
an absolute monarch, off duty a kindly citizen, and it seemed to all of
us who knew the country that he would die as he had lived, still in
harness. One did not expect too much—the so-called elections were
a pleasant farce, but the country was a deal better governed than the
western half of the United States. Any fellow entitled to a linen collar
in Europe wore a revolver in Mexico, as part of the dress of a
gentleman, but in the wildest districts I never carried a cartridge.
Diaz had made his country a land of peace and order, strong,
respected, prosperous, with every outward sign of coming greatness.
Excepting only Napoleon and the late Japanese emperor, he was
both in war and peace the greatest leader our world has ever known.
But the people proved unworthy of their chief; to-day he is a broken
exile, and Mexico has lapsed back into anarchy.
XIX
A. D. 1870
THE SPECIAL CORRESPONDENT
A LADY who remembers John Rowlands at the workhouse school

in Denbigh tells me that he was a lazy disagreeable boy. He is also
described as a “full-faced, stubborn, self-willed, round-headed,
uncompromising, deep fellow. He was particularly strong in the trunk,
but not very smart or elegant about the legs, which were
disproportionately short. His temperament was unusually secretive;
he could stand no chaff nor the least bit of humor.”
Perhaps that is why he ran away to sea; but anyway a sailing
ship landed him in New Orleans, where a rich merchant adopted him
as a son. Of course a workhouse boy has nothing to be patriotic
about, so it was quite natural that this Welsh youth should become a
good American, also that he should give up the name his mother
bore, taking that of his benefactor, Henry M. Stanley. The old man
died, leaving him nothing, and for two years there is no record until
the American Civil War gave him a chance of proving his patriotism
to his adopted country. He was so tremendously patriotic that he
served on both sides, first in the confederate army, then in the
federal navy. He proved a very brave man, and after the war,
distinguished himself as a special correspondent during an Indian
campaign in the West. Then he joined the staff of the New York
Herald serving in the Abyssinian War, and the civil war in Spain. He
allowed the Herald to contradict a rumor that he was a Welshman.
“Mr. Stanley,” said the paper, “is neither an Ap-Jones, nor an Ap-
Thomas. Missouri and not Wales is his birthplace.”
Privately he spent his holidays with his mother and family in
Wales, speaking Welsh no doubt with a strong American accent. The
whitewashed American has always a piercing twang, even if he has
adopted as his “native” land, soft-voiced Missouri, or polished
Louisiana.
In those days Doctor Livingstone was missing. The gentle daring
explorer had found Lakes Nyassa and Tanganyika, and to the
westward of them, a mile wide river, the Lualaba, which he supposed
to be headwaters of the Nile. He was slowly dying of fever, almost
penniless, and always when he reached the verge of some new
discovery, his cowardly negro carriers revolted, or ran away, leaving
him to his fate. No word of him had reached the world for years.
England was anxious as to the fate of one of her greatest men, so
there were various attempts to send relief, delayed by the expense,
and not perhaps handled by really first-rate men. To find Livingstone
would be a most tremendous world-wide advertisement, say for a
patent-pill man, a soap manufacturer, or a newspaper. All that was
needed was unlimited cash, and the services of a first-rate practical
traveler, vulgar enough to use the lost hero as so much “copy” for his
newspaper. The New York Herald had the money, and in Stanley, the
very man for the job.
Not that the Herald, or Stanley cared twopence about the fate of
Livingstone. The journal sent the man to make a big journey through
Asia Minor and Persia on his way to Zanzibar. The more
Livingstone’s rescue was delayed the better the “ad” for Stanley and
the Herald.
As to the journey, Stanley’s story has been amply advertised,
and we have no other version because his white followers died. He
found Livingstone at Ujiji on Lake Tanganyika, and had the grace to
reverence, comfort and succor a dying man.
As to Stanley’s magnificent feat of exploring the great lakes, and
descending Livingstone’s river to the mouth of the Congo, again his
story is well exploited while the version of his white followers is
missing, because they gave their lives.
In Stanley’s expedition which founded the Congo State, and in
his relief of Emin Pasha, the white men were more fortunate, and
some lived. It is rumored that they did not like Mr. Stanley, but his
negro followers most certainly adored him, serving in one journey
after another. There can be no doubt too, that with the unlimited
funds that financed and his own fine merits as a traveler, Stanley did
more than any other explorer to open up the dark continent, and to
solve its age-long mysteries. It was not his fault that Livingstone
stayed on in the wilderness to die, that the Congo Free State
became the biggest scandal of modern times, or that Emin Pasha
flatly refused to be rescued from governing the Soudan.
Henry M. Stanley
Stanley lived to reap the rewards of his great deeds, to forget

that he was a native of Missouri and a freeborn American citizen, to
accept the honor of knighthood and to sit in the British parliament.
Whether as a Welshman, or an American, a confederate, or a
federal, a Belgian subject or a Britisher, he always knew on which
side his bread was buttered.
XX
A. D. 1871
LORD STRATHCONA
IT is nearly a century now since Lord Strathcona was born in a

Highland cottage. His father, Alexander Smith, kept a little shop at
Forres, in Elgin; his mother, Barbara Stewart, knew while she reared
the lad that the world would hear of him. His school, founded by a
returned adventurer, was one which sent out settlers for the colonies,
soldiers for the army, miners for the gold-fields, bankers for England,
men to every corner of the world. As the lad grew, he saw the
soldiers, the sailors, the adventurers, who from time to time came
tired home to Forres, and among these was his uncle, John Stewart,
famous in the annals of the Canadian frontier, rich, distinguished,
commending all youngsters to do as he had done. When Donald
Smith was in his eighteenth year, this uncle procured him a clerkship
in the Hudson’s Bay Company.
Canada was in revolt when in 1837 the youngster reached
Montreal, for Robert Nelson had proclaimed a Canadian republic and
the British troops were busy driving the republicans into the United
States. So there was bloodshed, the burning of houses, the filling of
the jails with rebels to be convicted presently and hanged. Out of all
this noise and confusion, Donald Smith was sent into the silence of
Labrador, the unknown wilderness of the Northeast Territory, where
the first explorer, McLean, was searching for tribes of Eskimos that
might be induced to trade with the Hudson’s Bay Company. “In
September (1838),” wrote McLean, “I was gratified by the arrival of
despatches from Canada by a young clerk appointed to the district.
By him we received the first intelligence of the stirring events which
had taken place in the colonies during the preceding year.” So Smith
had taken a year to carry the news of the Canadian revolt to that
remote camp of the explorers.
Henceforward, for many years there exists no public record of
Donald Smith’s career, and he has flatly refused to tell the story lest
he should appear to be advertising. His work consisted of trading
with the savages for skins, of commanding small outposts, healing
the sick, administering justice, bookkeeping, and of immense
journeys by canoe in summer, or cariole drawn by a team of dogs in
winter. The winter is arctic in that Northeast Territory, and a very
pleasant season between blizzards, but the summer is cursed with a
plague of insects, black flies by day, mosquitoes by night almost
beyond endurance. Like other men in the service of the company,
Mr. Smith had the usual adventures by flood and field, the peril of the
snow-storms, the wrecking of canoes. There is but one story extant.
His eyesight seemed to be failing, and after much pain he ventured
on a journey of many months to seek the help of a doctor in
Montreal. Sir George Simpson, governor of the company, met him in
the outskirts of the city.
“Well, young man,” he said, “why are you not at your post?”
“My eyes, sir; they got so bad, I’ve come to see a doctor.”
“And who gave you permission to leave your post?”
“No one, sir.” It would have taken a year to get permission, and
his need was urgent.
“Then, sir,” answered the governor, “if it’s a question between
your eyes, and your service in the Hudson’s Bay Company, you’ll
take my advice, and return this instant to your post.”
Without another word, without a glance toward the city this man
turned on his tracks, and set off to tramp a thousand miles back to
his duty.
The man who has learned to obey has learned to command, and
wherever Smith was stationed, the books were accurate, the trade
was profitable. He was not heard of save in the return of profits,
while step by step he rose to higher and higher command, until at
the age of forty-eight he was appointed governor of the Hudson’s
Bay Company, sovereign from the Atlantic to the Pacific, reigning
over a country nearly as large as Europe. To his predecessors this
had been the crowning of an ambitious life; to him, it was only the
beginning of his great career.
The Canadian colonies were then being welded into a nation and
the first act of the new Dominion government was to buy from the
Hudson’s Bay Company the whole of its enormous empire, two
thousand miles wide and nearly five thousand miles long. Never was
there such a sale of land, at such a price, for the cash payment
worked out at about two shillings per square mile. Two-thirds of the
money went to the sleeping partners of the company in England;
one-third—thanks to Mr. Smith’s persuasion—was granted to the
working officers in Rupert’s Land. Mr. Smith’s own share seems to
have been the little nest egg from which his fortune has hatched.
When the news of the great land sale reached the Red River of
the north, the people there broke out in revolt, set up a republic, and
installed Louis Riel as president at Fort Garry.
Naturally this did not meet the views of the Canadian
government, which had bought the country, or of the Hudson’s Bay
Company, which owned the stolen fort. Mr. Smith, governor of the
company, was sent at once as commissioner for the Canadian
government to restore the settlement to order. On his arrival the
rebel president promptly put him in jail, and openly threatened his
life. In this awkward situation, Mr. Smith contrived not only to stay
alive, but to conduct a public meeting, with President Riel acting as
his interpreter to the French half-breed rebels. The temperature at
this outdoor meeting was twenty degrees below zero, with a keen
wind, but in course of five hours’ debating, Mr. Smith so undermined
the rebel authority that from that time it began to collapse. Afterward,
although the rebels murdered one prisoner, and times were more
than exciting, Mr. Smith’s policy gradually sapped the rebellion, until,
when the present Lord Wolseley arrived with British troops, Riel and
his deluded half-breeds bolted. So, thanks to Mr. Smith, Fort Garry is
now Winnipeg, the central city of Canada, capital of her central
province, Manitoba.
But when Sir Donald Smith had resigned from the Hudson’s Bay
Company’s service, and became a politician, he schemed, with
unheard-of daring, for even greater ends. At his suggestion, the
Northwest Mounted Police was formed and sent out to take
possession of the Great Plains. That added a wheat field to Canada
which will very soon be able to feed the British empire. Next he
speculated with every dollar he could raise, on a rusty railway track,
which some American builders had abandoned because they were
bankrupt. He got the rail head into Winnipeg, and a large trade
opened with the United States. So began the boom that turned
Manitoba into a populous country, where the buffalo had ranged
before his coming. Now he was able to startle the Canadian
government with the warning that unless they hurried up with a
railway, binding the whole Dominion from ocean to ocean, all this
rich western country would drift into the United States. When the
government had failed in an attempt to build the impossible railway,
Sir Donald got Montreal financiers together, cousins and friends of
his own, staked every dollar he had, made them gamble as heavily,
and set to work on the biggest road ever constructed. The country to
be traversed was almost unexplored, almost uninhabited except by
savages, fourteen hundred miles of rock and forest, a thousand
miles of plains, six hundred miles of high alps.
The syndicate building the road consisted of merchants in a
provincial town not bigger then than Bristol, and when they met for
business it was to wonder vaguely where the month’s pay was to
come from for their men. They would part for the night to think, and
by morning, Donald Smith would say, “Well, here’s another million—
that ought to do for a bit.” On November seven, 1885, he drove the
last spike, the golden spike, that completed the Canadian Pacific
railway, and welded Canada into a living nation.
Since then Lord Strathcona has endowed a university and given
a big hospital to Montreal. At a cost of three hundred thousand
pounds he presented the famous regiment known as Strathcona’s
Horse, to the service of his country, and to-day, in his ninety-third
year is working hard as Canadian high commissioner in London.
XXI
A. D. 1872
THE SEA HUNTERS
THE Japanese have heroes and adventurers just as fine as our

own, most valiant and worthy knights. Unhappily I am too stupid to
remember their honorable names, to understand their motives, or to
make out exactly what they were playing at. It is rather a pity they
have to be left out, but at least we can deal with one very odd phase
of adventure in the Japan seas.
The daring seamen of old Japan used to think nothing of
crossing the Pacific to raid the American coast for slaves. But two or
three hundred years ago the reigning shogun made up his mind that
slaving was immoral. So he pronounced an edict by which the
builders of junks were forbidden to fill in their stern frame with the
usual panels. The junks were still good enough for coastwise trade
at home, but if they dared the swell of the outer ocean a following
sea would poop them and send them to the bottom. That put a stop
to the slave trade; but no king can prevent storms, and law or no law,
disabled junks were sometimes swept by the big black current and
the westerly gales right across the Pacific Ocean. The law made only
one difference, that the crippled junks never got back to Japan; and
if their castaway seamen reached America the native tribes enslaved
them. I find that during the first half of the nineteenth century the
average was one junk in forty-two months cast away on the coasts of
America.
Now let us turn to another effect of this strange law that disabled
Japanese shipping. Northward of Japan are the Kuril Islands in a
region of almost perpetual fog, bad storms and bitter cold, ice pack,
strong currents and tide rips, combed by the fanged reefs, with
plenty of earthquakes and eruptions to allay any sense of monotony.
The large and hairy natives are called the Ainu, who live by fishing,
and used to catch sea otter and fur seal. These furs found their way
via Japan to China, where sea-otter fur was part of the costly official
winter dress of the Chinese mandarins. As to the seal, their whiskers
are worth two shillings a set for cleaning opium pipes, and one part
of the carcass sells at a shilling a time for medicine, apart from the
worth of the fur.
Now the law that disabled the junks made it impossible for Japan
to do much trade in the Kurils, so that the Russians actually got there
first as colonists.
But no law disabled the Americans, and when the supply of sea
otter failed on the Californian coast in 1872 a schooner called the
Cygnet crossed the Pacific to the Kuril Islands. There the sea-otters
were plentiful in the kelp beds, tame as cats and eager to inspect the
hunters’ boats. Their skins fetched from eighty to ninety dollars.
When news came to Japan of this new way of getting rich, a
young Englishman, Mr. H. J. Snow, bought a schooner, a hog-
backed relic called the Swallow in which he set out for the hunting.
Three days out, a gale dismasted her, and putting in for shelter she
was cast away in the Kuriles. Mr. Snow’s second venture was
likewise cast away on a desert isle, where the crew wintered. “My
vessels,” he says, “were appropriately named. The Swallow
swallowed up part of my finances, and the Snowdrop caused me to
drop the rest.”
During the winter another crew of white men were in quarters on
a distant headland of the same Island Yeturup, and were cooking
their Christmas dinner when they met with an accident. A dispute
had arisen between two rival cooks as to how to fry fritters, and
during the argument a pan of boiling fat capsized into the stove and
caught alight. The men escaped through the flames half dressed,
their clothes on fire, into the snow-clad wilderness and a shrewd
wind. Then they set up a shelter of driftwood with the burning ruin in
front to keep them warm, while they gravely debated as to whether
they ought to cremate the cooks upon the ashes of their home and of
their Christmas dinner.
To understand the adventures of the sea hunters we must follow
the story of the leased islands. The Alaska Commercial Company, of
San Francisco, leased the best islands for seal and otter fishing.
From the United States the company leased the Pribilof Islands in
Bering Sea, a great fur-seal metropolis with a population of nearly
four millions. They had armed native gamekeepers and the help of
an American gunboat. From Russia the company leased Bering and
Copper Islands off Kamchatka, and Cape Patience on Saghalian
with its outlier Robber Island. There also they had native
gamekeepers, a patrol ship, and the help of Russian troops and
gunboats. The company had likewise tame newspapers to preach
about the wickedness of the sea hunters and call them bad names.
As a rule the sea hunters did their hunting far out at sea where it was
perfectly lawful. At the worst they landed on the forbidden islands as
poachers. The real difference between the two parties was that the
sea hunters took all the risks, while the company had no risks and
took all the profits.
In 1883 Snow made his first raid on Bering Island. Night fell
while his crew were busy clubbing seals, and they had killed about
six hundred when the garrison rushed them. Of course the hunters
made haste to the boats, but Captain Snow missed his men who
should have followed him, and as hundreds of seals were taking to
the water he joined them until an outlying rock gave shelter behind
which he squatted down, waist-deep. When the landscape became
more peaceful he set off along the shore of boulders, stumbling,
falling and molested by yapping foxes. He had to throw rocks to keep
them off. When he found the going too bad he took to the hills, but
sea boots reaching to the hips are not comfy for long walks, and
when he pulled them off he found how surprisingly sharp are the
stones in an Arctic tundra. He pulled them on again, and after a long
time came abreast of his schooner, where he found one of the
seamen. They hailed and a boat took them on board, where the
shipkeeper was found to be drunk, and the Japanese bos’n much in
need of a thrashing. Captain Snow supplied what was needed to the
bos’n and had a big supper, but could not sail as the second mate
was still missing. He turned in for a night’s rest.
Next morning bright and early came a company’s steamer with a
Russian officer and two soldiers who searched the schooner. There
was not a trace of evidence on board, but on general principles the
vessel was seized and condemned, all her people suffering some
months of imprisonment at Vladivostok.
In 1888, somewhat prejudiced against the virtuous company,
Captain Snow came with the famous schooner Nemo, back to the
scene of his misadventure. One morning with three boats he went
prospecting for otter close along shore, shot four, and his hunters
one, then gave the signal of return to the schooner. At that moment
two shots rang out from behind the boulders ashore, and a third,
which peeled some skin from his hand, followed by a fusillade like a
hail storm. Of the Japanese seamen in Snow’s boat the boat steerer
was shot through the backbone. A second man was hit first in one
leg, then in the other, but went on pulling. The stroke oar, shot in the
calf, fell and lay, seemingly dead, but really cautious. Then the other
two men bent down and Snow was shot in the leg.
So rapid was the firing that the guns ashore must have heated
partly melting the leaden bullets, for on board the boat there was a
distinct perfume of molten lead. Three of the bullets which struck the
captain seem to have been deflected by his woolen jersey, and one
which got through happened to strike a fold. It had been noted in the
Franco-Prussian War that woolen underclothes will sometimes turn
leaden bullets.
“I remember,” says Captain Snow, “weighing the chances ... of
swimming beside the boat, but decided that we should be just as
liable to be drowned as shot, as no one could stand the cold water
for long. For the greater part of the time I was vigorously plying my
paddle ... and only presenting the edge of my body, the left side, to
the enemy. This is how it was that the bullets which struck me all
entered my clothing on the left side. I expected every moment to be
shot through the body, and I could not help wondering how it would
feel.”
With three dying men, and three wounded, he got the sinking
boat under sail and brought her alongside the schooner.
Of course it was very good of the Alaska Commercial Company
to preserve the wild game of the islands, but even gamekeepers may
show excess of zeal when it comes to wholesale murder. To all of us
who were in that trade it is a matter of keen regret that the officers
ashore took such good cover. Their guards, and the Cossacks, were
kindly souls enough, ready and willing—in the absence of the
officers to sell skins to the raiders or even, after some refreshments,
to help in clubbing a few hundred seals. It was rather awkward,
though, for one of the schooners at Cape Patience when in the midst
of these festivities a gunboat came round the corner.
The American and the Japanese schooners were not always
quite good friends, and there is a queer story of a triangular duel
between three vessels, fought in a fog. Mr. Kipling had the Rhyme of
the Three Sealers, he told me, from Captain Lake in Yokohama. I
had it from the mate of one of the three schooners, The Stella. She
changed her name to Adele, and the mate became master, a little,
round, fox-faced Norseman, Hans Hansen, of Christiania. In 1884
the Adele was captured by an American gunboat and taken to San
Francisco. Hansen said that he and his men were marched through
the streets shackled, and great was the howl about pirates, but when
the case came up for trial the court had no jurisdiction, and the ship
was released. From that event dates the name “Yokohama Pirates,”
and Hansen’s nickname as the Flying Dutchman. Because at the
time of capture he had for once been a perfectly innocent deep sea
sealer, he swore everlasting war against the United States,
transferred his ship to the port of Victoria, British Columbia, and
would hoist by turns the British, Japanese, German, Norwegian or
even American flag, as suited his convenience.
Once when I asked him why not the Black Flag, he grinned,
remarking that them old-fashioned pirates had no business sense.
Year after year he raided the forbidden islands to subvert the
garrisons, rob warehouses, or plunder the rookeries, while gunboats
of four nations failed to effect his capture. In port he was a pattern of
innocent virtue, at sea his superb seamanship made him as hard to

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To my dear little girl, Alexandra.

To my students, who have inspired me far more than

To my dear wife, Evelyn, who encouraged me to participate

To Drexel, who taught me about perseverance in the

To Jack Dainty and Bob Jefferies, who taught me

CHAPTER 1 Introduction to Ecology 2

CHAPTER 2 Environmental Inﬂuences 28

CHAPTER 3 Ecological Energetics 44

CHAPTER 4 Nutrients and Their Cycling 70

CHAPTER 5 Population Ecology 94

CHAPTER 6 Behavioural Ecology 134

CHAPTER 7 Physiological Ecology 158

CHAPTER 8 Life Histories 212

CHAPTER 9 Community Ecology 248

CHAPTER 10 Disturbance and Succession 296

CHAPTER 11 Biomes and Ecozones 338

CHAPTER 12 Biodiversity 368

CHAPTER 13 Landscape Ecology 392

CHAPTER 14 Conservation of the Natural World 430

CHAPTER 15 Resource Ecology 478

CHAPTER 16 Paleoecology: Lessons from the Past 534

CHAPTER 17 Ecology and Society 566

NEL BRIEF CONTENTS v

Bill Freedman is an ecol- University Biology Chairs. Beyond academics, Bill is a

of Botany. He has taught in scapes of nearby Shield

NEL ABOUT THE AUTHORS xi

Toronto in Mississauga. environmental change, focusing primarily on lake

a cross-appointment to the agencies, including a 3M National Teaching Fellowship

life as an Instructor Emeritus.

Environmental Studies from

NEL ABOUT THE AUTHORS xiii

mission is to develop, lead,

A LADY who remembers John Rowlands at the workhouse school

Stanley lived to reap the rewards of his great deeds, to forget

IT is nearly a century now since Lord Strathcona was born in a

THE Japanese have heroes and adventurers just as fine as our

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