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Contents vii
8.2 Mate Choice and Resource Provisioning

Concept 8.2 Review 182
179
Chapter 10 Population Dynamics 218
Concepts 218
8.3 Nonrandom Mating in a Plant Population 182
Concept 8.3 Review 184 10.1 Dispersal 220
8.4 Sociality 184

Dispersal of Expanding Populations 220
Range Changes in Response to Climate Change 221
Cooperative Breeders 185
Dispersal in Response to Changing Food Supply 222
Investigating the Evidence 8: Estimating Heritability Using
Regression Analysis 188 Dispersal in Rivers and Streams 223
Concept 8.4 Review 191 Concept 10.1 Review 224
10.2 Metapopulations 224
8.5 Eusociality 191
Eusocial Species 191 A Metapopulation of an Alpine Butterfly 225
Evolution of Eusociality 193 Dispersal Within a Metapopulation of Lesser Kestrels 226
10.3 Patterns of Survival 227
Applications: Behavioral Ecology and Conservation 195
Estimating Patterns of Survival 227
Tinbergen’s Framework 195
High Survival Among the Young 227
Environmental Enrichment and Development
of Behavior 195 Constant Rates of Survival 229
High Mortality Among the Young 230
Three Types of Survivorship Curves 230
Section III 10.4 Age Distribution 231
Contrasting Tree Populations 231
POPULATION ECOLOGY A Dynamic Population in a Variable Climate 232
Chapter 9 Population Distribution 10.5 Rates of Population Change 233
and Abundance 198 Estimating Rates for an Annual Plant 233
Concepts 198 Estimating Rates When Generations Overlap 234
Investigating the Evidence 10: Hypotheses and Statistical
9.1 Distribution Limits 200
Significance 236
Kangaroo Distributions and Climate 200
A Tiger Beetle of Cold Climates 201
Applications: Changes in Species Distributions in Response
Distributions of Plants Along a Moisture-Temperature
to Climate Warming 237
Gradient 202
Distributions of Barnacles Along an Intertidal Exposure
Gradient 203 Chapter 11 Population Growth 241
Concept 9.1 Review 204 Concepts 241
9.2 Patterns on Small Scales 204 11.1 Geometric and Exponential Population
Scale, Distributions, and Mechanisms 205 Growth 242
Distributions of Tropical Bee Colonies 205 Geometric Growth 242
Distributions of Desert Shrubs 206 Exponential Growth 243
Concept 9.2 Review 208 Exponential Growth in Nature 244
9.3 Patterns on Large Scales 208 Concept 11.1 Review 245
Bird Populations Across North America 208 11.2 Logistic Population Growth 246
Investigating the Evidence 9: Clumped, Random, Concept 11.2 Review 248
and Regular Distributions 209 11.3 Limits to Population Growth 248
Plant Distributions Along Moisture Gradients 210 Environment and Birth and Death Among Darwin’s
Concept 9.3 Review 211 Finches 249
9.4 Organism Size and Population Density 212 Investigating the Evidence 11: Frequency of Alternative
Animal Size and Population Density 212 Phenotypes in a Population 250
Plant Size and Population Density 212 Concept 11.3 Review 253
Concept 9.4 Review 213 Applications: The Human Population 253
Applications: Rarity and Vulnerability Distribution and Abundance 253
to Extinction 214 Population Dynamics 254
Seven Forms of Rarity and One of Abundance 214 Population Growth 254
viii Contents
Chapter 12 Life Histories 258 Investigating the Evidence 13: Field Experiments
299
Concepts 258 Applications: Competition between Native

12.1 Offspring Number Versus Size 259 and Invasive Species 300
Egg Size and Number in Fish 260
Seed Size and Number in Plants 262
Seed Size and Seedling Performance 263
Chapter 14 Exploitative Interactions: Predation,
Concept 12.1 Review 265 Herbivory, Parasitism, and
12.2 Adult Survival and Reproductive Allocation 266 Disease 303
Life History Variation Among Species 266 Concepts 303
Life History Variation Within Species 267 14.1 Complex Interactions 304
Concept 12.2 Review 270 Parasites and Pathogens that Manipulate Host
12.3 Life History Classification 270 Behavior 304
r and K Selection 270 The Entangling of Exploitation with Competition 307
Plant Life Histories 271 Concept 14.1 Review 308
Investigating the Evidence 12: A Statistical Test 14.2 Exploitation and Abundance 308
for Distribution Pattern 272 A Herbivorous Stream Insect and Its Algal Food 308
Opportunistic, Equilibrium, and Periodic Life Bats, Birds, and Herbivory in a Tropical Forest 309
Histories 274
A Pathogenic Parasite, a Predator, and Its Prey 311
Lifetime Reproductive Effort and Relative Offspring Size:
Two Central Variables? 275
Concept 12.3 Review 276 14.3 Dynamics 312
Cycles of Abundance in Snowshoe Hares and Their
Applications: Climate Change and Timing of Reproduction
Predators 312
and Migration 277
Investigating the Evidence 14: Standard Error of the
Altered Plant Phenology 277 Mean 314
Animal Phenology 278 Experimental Test of Food and Predation Impacts 316
Population Cycles in Mathematical and Laboratory
Section IV Models 317

INTERACTIONS 14.4 Refuges 320
Refuges and Host Persistence in Laboratory
Chapter 13 Competition 282 and Mathematical Models 320
Exploited Organisms and Their Wide Variety
Concepts 282 of “Refuges” 321
13.1 Intraspecific Competition 284 Concept 14.4 Review 323
Intraspecific Competition Among Plants 284 14.5 Ratio-Dependent Models of Functional Response 323
Intraspecific Competition Among Planthoppers 285 Alternative Model for Trophic Ecology 324
Interference Competition Among Terrestrial Isopods 285 Evidence for Ratio-Dependent Predation 324
13.2 Competitive Exclusion and Niches 286 Applications: The Value of Pest Control by Bats:
The Feeding Niches of Darwin’s Finches 286 A Case Study 327
The Habitat Niche of a Salt Marsh Grass 288
13.3 Mathematical and Laboratory Models 289
Chapter 15 Mutualism 331
Concepts 331
Modeling Interspecific Competition 289
Laboratory Models of Competition 291 15.1 Plant Mutualisms 332
Concept 13.3 Review 292 Plant Performance and Mycorrhizal Fungi 333
13.4 Competition and Niches 292 Ants and Swollen Thorn Acacias 336
Niches and Competition Among Plants 293 A Temperate Plant Protection Mutualism 340
Niche Overlap and Competition between Barnacles 293 Concept 15.1 Review 341
Competition and the Habitat of a Salt Marsh Grass 295 15.2 Coral Mutualisms 341
Competition and the Niches of Small Rodents 295 Zooxanthellae and Corals 342
Character Displacement 296 A Coral Protection Mutualism 342
Evidence for Competition in Nature 298 Concept 15.2 Review 344
Contents ix
15.3 Evolution of Mutualism 344 17.2 Indirect Interactions 376

Investigating the Evidence 15: Confidence Intervals 345 Indirect Commensalism 376
Facultative Ant-Plant Protection Mutualisms 347 Apparent Competition 376
Applications: Mutualism and Humans 348 17.3 Keystone Species 378
Guiding Behavior 348 Food Web Structure and Species Diversity 379
Experimental Removal of Sea Stars 380
Snail Effects on Algal Diversity 381
Section V Fish as Keystone Species in River Food Webs 383
Investigating the Evidence 17: Using Confidence Intervals
COMMUNITIES AND ECOSYSTEMS to Compare Populations 384
Chapter 16 Species Abundance 17.4 Mutualistic Keystones 386
and Diversity 352 A Cleaner Fish as a Keystone Species 386
Concepts 352 Seed Dispersal Mutualists as Keystone Species 387
16.1 Species Abundance 354 Concept 17.4 Review 388
The Lognormal Distribution 354 Applications: Human Modification of Food Webs 388
Concept 16.1 Review 355 The Empty Forest: Hunters and Tropical Rain Forest
16.2 Species Diversity 355 Animal Communities 388
Ants and Agriculture: Keystone Predators for Pest
A Quantitative Index of Species Diversity 355
Control 389
Rank-Abundance Curves 356
16.3 Environmental Complexity 357 Chapter 18 Primary and Secondary
Forest Complexity and Bird Species Diversity 358 Production 392
Investigating the Evidence 16: Estimating the Number
of Species in Communities 359 Concepts 392
Niches, Heterogeneity, and the Diversity of Algae and 18.1 Patterns of Terrestrial Primary Production 394
Plants 360 Actual Evapotranspiration and Terrestrial Primary
The Niches of Algae and Terrestrial Plants 360 Production 394
Complexity in Plant Environments 361 Soil Fertility and Terrestrial Primary Production 395
Soil and Topographic Heterogeneity and the Diversity Concept 18.1 Review 396
of Tropical Forest Trees 361 18.2 Patterns of Aquatic Primary Production 396
Algal and Plant Species Diversity and Increased Nutrient
Patterns and Models 396
Availability 363
Whole Lake Experiments on Primary
Nitrogen Enrichment and Ectomycorrhizal Fungus
Production 397
Diversity 363
Global Patterns of Marine Primary Production 397
16.4 Disturbance and Diversity 364
18.3 Primary Producer Diversity 399
The Nature and Sources of Disturbance 364
Terrestrial Plant Diversity and Primary Production 399
The Intermediate Disturbance Hypothesis 364
Algal Diversity and Aquatic Primary Production 400
Disturbance and Diversity in the Intertidal Zone 365
Disturbance and Diversity in Temperate Grasslands 365
Concept 16.4 Review 367 18.4 Consumer Influences 401
Piscivores, Planktivores, and Lake Primary
Applications: Disturbance by Humans 367
Production 401
Urban Diversity 368 Grazing by Large Mammals and Primary Production
on the Serengeti 403
Chapter 17 Species Interactions Concept 18.4 Review 405
and Community Structure 372 18.5 Secondary Production 405
Investigating the Evidence 18: Comparing Two Populations
Concepts 372
with the t-Test 406
17.1 Community Webs 374 A Trophic Dynamic View of Ecosystems 406
Detailed Food Webs Reveal Great Complexity 374 Linking Primary Production
Strong Interactions and Food Web Structure 374 and Secondary Production 408
x Contents
Applications: Using Stable Isotope Analysis to Study Feeding Successional Mechanisms in the Rocky Intertidal
Habits 410 Zone 447
Using Stable Isotopes to Identify Sources of Energy Successional Mechanisms in Forests 449
in a Salt Marsh 410 Concept 20.3 Review 450
20.4 Community and Ecosystem Stability 450
Chapter 19 Nutrient Cycling Lessons from the Park Grass Experiment 451
and Retention 414 Replicate Disturbances and Desert Stream Stability 451
Concepts 414
Investigating the Evidence 20: Variation Around the
19.1 Nutrient Cycles 415 Median 454
The Phosphorus Cycle 416 Applications: Ecological Succession Informing Ecological
The Nitrogen Cycle 417 Restoration 454
The Carbon Cycle 418 Applying Succession Concepts to Restoration 455
19.2 Rates of Decomposition 419
Decomposition in Two Mediterranean Woodland
Ecosystems 419
Section VI
Decomposition in Two Temperate Forest Ecosystems 420 LARGE-SCALE ECOLOGY
Decomposition in Aquatic Ecosystems 422
Investigating the Evidence 19: Assumptions for Statistical Chapter 21 Landscape Ecology 460
Tests 423
Concepts 460
19.3 Organisms and Nutrients 425 21.1 Landscape Structure 462
Nutrient Cycling in Streams and Lakes 425 The Structure of Six Landscapes in Ohio 462
Animals and Nutrient Cycling in Terrestrial The Fractal Geometry of Landscapes 464
Ecosystems 427 Concept 21.1 Review 465
Plants and the Nutrient Dynamics of Ecosystems 428 21.2 Landscape Processes 465
Concept 19.3 Review 429 Landscape Structure and the Dispersal of Mammals 466
19.4 Disturbance and Nutrients 429 Habitat Patch Size and Isolation and the Density
Disturbance and Nutrient Loss from Forests 429 of Butterfly Populations 467
Flooding and Nutrient Export by Streams 430 Habitat Corridors and Movement of Organisms 468
Concept 19.4 Review 431 Landscape Position and Lake Chemistry 469
Investigating the Evidence 21: Comparison of Two Samples
Applications: Altering Aquatic and Terrestrial
Using a Rank Sum Test 470
Ecosystems 432
Chapter 20 Succession and Stability 435

21.3 Origins of Landscape Structure and Change
Geological Processes, Climate, and Landscape
471
Concepts 435 Structure 472

20.1 Community Changes During Succession 437 Organisms and Landscape Structure 474
Primary Succession at Glacier Bay 437 Fire and the Structure of a Mediterranean Landscape 478
Secondary Succession in Temperate Forests 438 Concept 21.3 Review 479
Succession in Rocky Intertidal Communities 439 Applications: Restoring a Riverine Landscape 479
Succession in Stream Communities 439 Riverine Restoration: The Kissimmee River 479
20.2 Ecosystem Changes During Succession 440 Chapter 22 Geographic Ecology 484
Ecosystem Changes at Glacier Bay 441
Concepts 484
Four Million Years of Ecosystem Change 441
Recovery of Nutrient Retention 22.1 Area, Isolation, and Species Richness 486
Following Disturbance 443 Island Area and Species Richness 486
Succession and Stream Ecosystem Properties 445 Island Isolation and Species Richness 488
20.3 Mechanisms of Succession 446 22.2 The Equilibrium Model of Island Biogeography 489
Facilitation 446 Species Turnover on Islands 490
Tolerance 446 Experimental Island Biogeography 491
Inhibition 446 Colonization of New Islands by Plants 492
Contents xi
Manipulating Island Area 493 El Niño and Marine Populations 511

Island Biogeography Update 494 El Niño and the Great Salt Lake 513
Concept 22.2 Review 494 El Niño and Terrestrial Populations in Australia 513
22.3 Latitudinal Gradients in Species Richness 494 Concept 23.1 Review 515
Latitudinal Gradient Hypotheses 494 23.2 Human Activity and the Global Nitrogen
Area and Latitudinal Gradients in Species Richness 496 Cycle 515
Continental Area and Species Richness 497 Concept 23.2 Review 516
Concept 22.3 Review 498 23.3 Changes in Land Cover 516
22.4 Historical and Regional Influences 498 Tropical Deforestation 516
Exceptional Patterns of Diversity 498 Concept 23.3 Review 519
Investigating the Evidence 22: Sample Size Investigating the Evidence 23: Discovering What’s Been
Revisited 499 Discovered 520
Historical and Regional Explanations 500 23.4 Human Influence on Atmospheric
Concept 22.4 Review 501 Composition 520
Applications: Global Positioning Systems, Remote Sensing, Depletion and Recovery of the Ozone Layer 523
and Geographic Information Systems 501 Concept 23.4 Review 524
Global Positioning Systems 502 Applications: Impacts of Global Climate Change 525
Remote Sensing 502 Shifts in Biodiversity and Widespread Extinction
Geographic Information Systems 503 of Species 525
Human Impacts of Climate Change 526
Chapter 23 Global Ecology 506 Appendix Statistical Tables 529
Concepts 506 Glossary 533
The Atmospheric Envelope and the Greenhouse Earth 507
References 543
23.1 A Global System 508
Photo Credits 554
The Historical Thread 509
El Niño and La Niña 510 Index 555
C
Contents
on
onte
tenntts xiii
x
xi
iiiii
Preface
This book was written for students taking their first under- to engage students and draw them into the discussion that
graduate course in ecology. I have assumed that students follows.
in this one-semester course have some knowledge of basic
Concepts: The goal of this book is to build a foundation of
chemistry and mathematics and have had a course in general
ecological knowledge around key concepts. I have found that
biology, which included introductions to physiology, biologi-
while beginning ecology students can absorb a few central
cal diversity, and evolution.
concepts well, they can easily get lost in a sea of details. The
key concepts are listed at the beginning of each chapter to
Organization of the Book
alert the student to the major topics to follow and to provide a
An evolutionary perspective forms the foundation of the place where the student can find a list of the important points
entire textbook, as it is needed to support understanding covered in each chapter. The sections in which concepts are
of major concepts. The textbook begins with a brief intro- discussed focus on published studies and, wherever possible,
duction to the nature and history of the discipline of ecol- the scientists who did the research are introduced. This case-
ogy, followed by section I, which includes two chapters on study approach supports the concepts with evidence, and
natural history—life on land and life in water and a chapter introduces students to the methods and people that have cre-
on population genetics and natural selection. Sections II ated the discipline of ecology. Each concept discussion ends
through VI build a hierarchical perspective through the with a series of concept review questions to help students
traditional subdisciplines of ecology: section II concerns test their knowledge and to reinforce key points made in the
adaptations to the environment; section III focuses on discussion.
population ecology; section IV presents the ecology of
interactions; section V summarizes community and ecosys-
Confirming Pag
tem ecology; and finally, section VI discusses large-scale es
ecology and includes chapters on landscape, geographic,

and global ecology. These topics were first introduced in
SEC TIO N
II Adaptations
to the Environ
ment
section I within a natural history context. In summary, the

book begins with the natural history of the planet, consid-
ers portions of the whole in the middle chapters, and ends
with another perspective of the entire planet in the con-
cluding chapter. The features of this textbook were care-
fully planned to enhance the students’ comprehension of
the broad discipline of ecology. Temperature
Relations
5
Features Designed with the
Student in Mind A group of Japa
conserving their
ity to regulate
nese macaques
body heat in the
, Macaca fusca
ta, huddles toge
midst of driving ther,
body snow. The capa
physiological adap temperature, using behaviora c-
tations, enables l, anatomical, and
cold winters in these monkeys
All chapters are based on a distinctive learning system, fea- Nagano, Japan,
site of the 1998
to live through
Winter Olympics.
the
in environmen
regulating bod
tal temperatu
y temperature.
re by
Concept 5.4 Rev 109
iew 119
turing the following key components: CHAPTER CO
NCEPTS 5.5 Many orga
nisms survive
temperatures extreme
by entering a
5.1 Macroclima stage. 119 resting
Student Learning Outcomes: Educators are being asked landscape to
te inte
variation in tem
racts with the
produce microcl
ima
local Concept 5.5 Rev
iew 121
perature. 100 tic Applications:
increasingly to develop concrete student learning outcomes Concept 5.1 Rev
iew 103
Local Extinct
an Urban Hea ion of a Land
t Island 122 Snail in
5.2 Adapting Summary 123
to one
for courses across the curriculum. In response to this need conditions gen set of environmental
a population’
erally reduce
s
Key Terms
124
s fitness in oth Review Questio
er
and to help focus student progress through the content, all environments.
Concept 5.2 Rev
103
iew 104
ns 124
sections of each chapter in the seventh edition begin with a 5.3 Most species
narrow range
perform best
of temperatu
in a fairly
LEARNING OU
TCOMES
res. 105 After studying
this section you
list of detailed student learning outcomes. Investigating
the Evidence
Laboratory Exp 5: 5.1 Distinguish betw
should be able
een temperature
to do
the following:
eriments 106 5.2 Explain the ecol and heat.
Concept 5.3 Rev ogical significa
iew 109 tal temperature
s.
nce of environm
en-
Introduction: The introduction to each chapter presents 5.4 Many orga
nisms have evo
T
ways to compen lved he thermometer
sate for variatio was one of the
the student with the flavor of the subject and important ns appear in the scie
suring and repo
ntific tool kit and
first instruments
rting temperature we have been mea-
to
what do thermom s ever since. How
background information. Some introductions include eters actually
quantify? Tem ever,
perature is a
historical events related to the subject; others pre- moL37282_ch05_

99
099-124.indd
sent an example of an ecological process. All attempt 99
29/09/14 9:13
pm xiii
xiv Preface
Illustrations: A great deal of effort has been put into the devel- expressions that arise to help students overcome these chal-
opment of illustrations, both photographs and line art. The goal lenges. In some cases, mathematical expressions are dissected
has been to create more effective pedagogical tools through in illustrations designed to complement their presentation in
skillful design and use of color, and to rearrange the traditional the associated narrative.
presentation of information in figures and
captions. Much explanatory material is lerian (honeybee
) and (b) nonpoi
sonous Batesia
n (hoverfly) mim
located within the illustrations, providing ic.
students with key information where they Birds leave the

population dom
need it most. The approach also provides by better camouf inated
laged individual
s.
an ongoing tutorial on graph interpreta-
tion, a skill with which many introductory
students need practice.
Detailed Explanations of Mathematics:
The mathematical aspects of ecology
commonly challenge many students
taking their first ecology course. This Birds eat a disp
roportionate num
of the conspicuou ber
text carefully explains all mathematical peppered moth
s members of a
population.
Figure 7.16
Birds and other
pre dators act as age
nts of natural sele
ction for improv
ed prey defens
of these, a mo e.
th and a fly. He
bald-faced horne inrich
ts have a prey cap ’s observations indicate
Though elusiv ture rate of less While some of
e the than 1% the items th t
G
Visualizing a process involving a predator and its prey.
To allow comparisons to
other Subtracting number of death
studies, number of Dall s
sheep from number alive at the
surviving and dying withi
n each beginning of each year gives
year of life is converted
to the number alive at the
numbers per 1,000 births
. beginning of the next year.
Number of
Age (years) survivors Number of deaths
at beginning during year
of year
0–1 1,000 199 and Ecosystems
1–2 801 1,000–199
12
2–3 789 801–12 13 By reducing planktivorous
3–4 776 789–13 fish
4–5 764
12 populations, piscivores indir
ectly t
5–6
30 increase populations of large a
734 46 zooplankton and indirectly
6–7 etc. moL37282_ch0 reduce
688 48
7_149-172.indd
161 biomass of phytoplankto
7–8 n.
640 69 Lake food web t
8–9 571
9–10
132 (
439 187
10–11 252
27/08/14 10:54 pm
le
156
11–12 96 p
90
12–13 6 3
b
13–14 Piscivores
3 3 sm
14–15 0 pl
Planktivorous fish sp
Plotting age on the x-axi
s
Planktivorous
and number of survivors invertebrates pl
ary production
of on the y-axis creates a tio

survivorship curve.
pi- Dall sheep surviving their

first year Large herbivorous
log
of life have a high proba Small herbivorous
le, surviving to about age 9.
bility of zooplankton zooplankton
led
prim
ge the
ng 1,000
in
Top-down influences on
to De
ly Sheep 10 years
old and older are So,
r-
easier prey for fed
Number of survivors
100 wolves and die

ll at a high rate. Large phytoplankton
with
of Survivorship curves are Small phytoplankton cal
plotted using a log
e 10
scale on the y-axis. man
s 10 large
n
h
large
e
t of p
1 at th
0 2 4 6 8 10 12 14 ton
Age (years)
Nutrients
Figure 10.14 Dall sheep Fig ure 18.12 The trophic casc zoop
: from life table to survi “cascading” indirect inter ade hypothesis, a result mary
(data from Murie 1944). vorship curve
actions. of
years is l Th
Helps students work with and interpret quantitative informa- Provides a visual representation of a hypothesis involving a
tion, involving converting numerical information into a graph. set of complex ecological interactions.
Preface xv
“Investigating the Evidence” Boxes: These readings offer chapter is organized are boldfaced and redefined in the
“mini-lessons” on the scientific method, emphasizing statis- summary to reemphasize the main points of the chapter.
tics and study design. They are intended to present a broad • Key Terms The listing of key terms provides page num-
outline of the process of science, while also providing step- bers for easy reference in each chapter.
by-step explanations. The series of boxes begins in chapter 1 • Review Questions The review questions are designed
with an overview of the scientific method, which establishes to help students think more deeply about each concept
a conceptual context for more specific material in the next and to reflect on alternative views. They also provide
21 chapters. The last reading wraps up the series with a dis- a place to fill in any remaining gaps in the information
cussion of electronic literature searches. Each Evidence box presented and take students beyond the foundation estab-
ends with one or more questions, under the heading “Critiqu- lished in the main body of the chapter.
ing the Evidence.” This feature is intended to stimulate criti-
End-of-Book Material:
cal thinking about the box content.
• Appendixes One appendix, “Statistical Tables,” is
Applications: Many undergraduate students want to know available to the student for reference. Answers to Con-
how abstract ideas and general relationships can be applied to cept Review questions and answers to Critiquing the
the ecological problems we face in the contemporary world. Evidence are now available with the book’s instructor
They are concerned with the practical side of ecology and resources.
want to know more about how the tools of science can be • Glossary List of all key terms and their definitions.
applied. Including a discussion of applications in each chapter • References References are an important part of any
motivates students to learn more of the underlying principles scientific work. However, many undergraduates are dis-
of ecology. In addition, it seems that environmental problems tracted by a large number of references within the text.
are now so numerous and so pressing that they have erased a One of the goals of a general ecology course should be to
once easy distinction between general and applied ecology. introduce these students to the primary literature without
burying them in citations. The number of citations has
End-of-Chapter Material:
been reduced to those necessary to support detailed dis-
• Summary The chapter summary reviews the main cussions of particular research projects.
points of the content. The concepts around which each • Index
Confirming Pag Confirming Pag

es es
106 122
Section II Section II
Adaptations to Adaptations to
the Environmen the Environmen
Applications
t t
Investigating
the Evidence 5 find the snail at
16 sites. Eight
ized, which mad of thes
Laboratory Ex Local Extinctio e the habitat unsu e sites had been urban-
periments n of a because natural itable for any
in an Urban He Land Snail
vege land
and 1990 the urba tation had been removed. Betw snails
at Island However, the eigh
nized area of Bas
el had increase
een 1900
LEARNING OU t other sites whe d by 500%.
LEARNING OU TCOMES appeared were re A. arbustorum
TCOMES After studying this section still covered by
vege had dis-
After studying you should be able . Four of these tation that app
this section you from both populat able to do the follo sites were cove eared suit-
should be able
to do the following ions used in the 5.21 Outline wing: three were on red by deciduo
5.12 Describ : body mass of experiments had changes in the
distribution of
riverbanks, and
one was on a railw us forest,
e the basic desi approximately an average Arianta arbustor the ment. These vege
5.13 Discuss gn of a laborato
ry experiment. may differ phy 5.4 g. Since mal um around Bas snai l tate d sites also supp ay emb ank-
the relative stre siologically, Ang es and females between 1900 el, Switzerland other land snai orted populations
ngths and weakne illetta included and 1990. , l species, of five
laboratory expe
riments and field sses of equal numbers
of males and fem approximately 5.22 Explain
how urbanization What caused the including C. nemoralis.
ecological stud observations in also was careful ales in his expe generally creates still supported extinction of
ies.
of light and to
to expose all the
lizards to the sam
riments. He island.” a “heat other snails? The A. arbustorum at sites that
the same numbers e quality 5.23 Review teristics of thes Baurs compare
ness and he mai of hours of ligh the evidence that e sites with thos d the charac-
One of the mos ntained them in t and dark- around the city temperature chan torum had pers e of the sites whe
t powerful way enclosures. Ang the same kinds of Basel are resp ges isted. They foun re A. arbus-
an experiment. s to test a hypothe illetta also fed of experimental extinctions of the onsible for loca two groups of d no difference
Experiments used sis is through the same type all the lizards in snail Arianta arbu l sites in regard between these
into one of two by ecologists gen of food: live cric his experiment storum. height of vege to slope, percent
categories—field erally fall these are the maj kets. tation, distance plant cover,
tory experiments.
Field and labo
experiments and
labora- or factors controlle The list could go on but Between 1906
and 1908, a Ph.D
land snail spec
ies present. The
from water, or
number of othe
provide complem ratory experime Now, what fact d in this experime (1909) studied . candidate nam uncovered was first major diffe r
entary informa nts generally ors did Angillet nt. land snails in the ed G. Bollinger in altitude. The rence the Baurs
somewhat in tion or evidence For each study ta vary in that Eighty-five year vicinity of Bas extinct had an sites where A.
their design. Her , and differ population, New experiment? s later, Bruno and el, Switzerland average altitude arbustorum was
laboratory expe e we discuss varied a single Jersey or South resurveyed Boll Anette Baur (199 . survived had an of 274 m. The
riments. the design of factor: tempera Carolina, he inger’s study sites 3) carefully aver plac es where it
letta maintained ture. In the expe
riment, Angil- land snails. In the near the snail had surv age altitude of 420 m. The plac
In a laboratory
experiment, the three temperature
lizards from New
Jersey and Sou process, they foun Basel for the presence of ived were also es where
all factors relative researcher attem s: 308, 338, and th Carolina at cies, Arianta arbu d that at least one A thermal ima cooler.
ly pts to keep 368C and estim storum, had disa snail spe- ge of the landscap
not kept constant constant except one. The one of metabolizab
le energy inta ated their rates sites. This disc ppeared from seve showed that surf e taken from a
is the one of inte factor that is overy led the Bau ral ace temperature
it is the one that rest to the expe Angilletta’s expe
riment revealed
ke at these thre
e temperatures. that may have prod rs to explore the of the ranged from abo s in summer arou satellite
the experimenter rimenter and tions have a max that lizards from uced extinction mechanisms ut 178 to 32.58C. nd Basel
conditions. Let’ varies across expe of thes A. arbustorum Surf
s draw an exam rimental This result sugg
imum metabol
izable energy inta
both popula- A. arbustorum
is a common land e local populations. had survived aver ace temperatures where
discussed in this ple of a laborato ests, ke at 338C. ests, and other snail in mea while the sites aged approximat
studies, Michael
chapter (see p.
000). Based upo
ry experiment optimum tempera contrary to the study’s hypothe central Europe
moist, vegetate
d habitats in nort dows, for- temperatures that
where the spec
ies had gone exti ely 228C,
Angilletta (200 n published ture for feeding sis, that the . The species hwestern and averaged approxi nct had surface
cally separated 1) concluded that populations. How does not differ in the Alps. The live s at altitudes up where the snai mately 258C.
populations of geog ever, the experime for the two Baurs report that to 2,700 m l was extinct The sites
porus undulatus, the eastern fenc raphi- S. undulatus from nt also showed at 2 to 4 years the snail is sexu hot areas with were
may differ phy e lizard, Scelo- energy intake com
South Carolina that
have a higher met at 338C and may live up ally mature temperatures grea also much closer to very
Angilletta desi siologically or pared to lizards abolizable shell diameter to 14 years. Adu based on the Bau ter than 298C.
gned a laborato behaviorally. provides evidence from New Jersey. s of 16 to 20 lt snails have rs’ thermal ima Figure 5.34 is
hypothesis that ry experiment of This result ditic. Though indi mm. The spec and shows whe ge of the area
ies is hermaph
significantly diffe
populations of
S. undulatus from to test the thought might exis the geographic differences that torum, they can
viduals general
ly mate with othe ro- The Baurs attri
re the snail was
extinct and whe
around Basel
rent climates regions with t across the rang Angilletta fertilize their own r A. arbus- buted the high re it persisted.
affects their rate differ in how of this experime e of S. undulatus. to three batches eggs. Adults prod sites where the er temperature
s of metabolizab temperature nt to reveal the The power of 20 to 80 eggs uce one snail s at the eight
of that experime le energy intake. ard performance
resulted from the
influence of tem
perature on liz- eggs in moss, each year. The
y deposit their from the urbanize is extinct to heating by thermal
nt are summar The results control all sign under plant litte d area s of the city. Bui radiation
want to consider ized by figure
5.10 ificant factors but ability of the researcher to hatch in 2 to 4
weeks, depend
r, or in the soil
. Egg store more heat
than vegetation. ldings and pave
duced those resu
here is the desi
gn of the experime What we
. the main factor
of interest was
the one of interest.
In this case is an especially ing upon tempera s generally of evaporation In addition, the ment
lts. What factors nt that pro- temperature. sensitive stage ture. The egg from vegetatio cooling effect
have attempted do you think Ang A. arbustorum in the life cycl over . Incr eased heat stor n is lost whe n an area is buil
to control in this illet often lives alon e of land snails.
similar numbers experiment? Firs ta may CRITIQUING THE
EVIDENCE 5 snail with a broa gside Cepea nem
oralis, a land
ized landscapes age and reduced
cooling make urba
t
of lizards from t, he used der geographic thermal islands. n-
20 lizards from the two populat 1. What is the southern Scandin distribution that centers is tran Heat energy stor
ions. He tested extends from sferred to the
at 308 and 368C,
both populations
at 338C, 13 from ecological rese
greatest strength
of laboratory expe How did the
avia to the Iber
ian peninsula. thermal radiatio surrounding land ed in urban
and New Jersey arch? riments in n, H. scap e through
second factor that 14 from South Carolina at 308 2. Why do ecol A. arbustorum? Baurs docume
nt local extincti The Baurs doc r
Angilletta cont and 368C. A ogists generally If you think abo ons of umented higher
rolled was lizar resulting from supplement info realize that it is ut it a bit, you near Basel whe temperatures at
d size. Lizards laboratory expe rmation usually easier will probably re A. arbustor the sites
the United Stat tions or experime rime nts with field obse species than its to determine the well-studied mec um is extinct
es, living in a broa nts? rva- absence. If you presence of a han ism and identified
(fig. 5.9). Taking d diversity of clim ing a survey, it do not encounte peratures of thes that could produce a
may be that you r a species dur- e sites. However the high
conditions, Mic
advantage of this
wide range of envi
atic zones
He collected a Fortunately, the just didn’t look ences they obse , are the tempera er tem-
hael Angilletta ronmental sample of liza Baurs had over hard enough. rved sufficient ture differ-
relations of S. (2001) studied maintained port rds from both fieldwork on A. 13 years of expe the warmer sites to exclude A. arbu
undulatus over the temperature ions of his sam populations and arbu rience doing ? The research storum from
his studies, Ang a portion of its 308, 338, and 36 ples from both For instance, they storum and knew its natural relations of A. ers compared the
illetta determin range. In one of 8C. Angilletta populations at knew that it is history well. arbustorum and
C. nemoralis
temperature
metabolizable ed how temperature rate enclosures kept his study after rainstorms, best to search clues. They con to
lizards in sepa for the snails centrated their
amount of ener
energy intake,
or ME influ ence s weighed to the
and prov ided them with cric - active. Consequ
when up to 70%
of the adult pop perature on repr studies on the influ find some
gy consumed (C) I. He measured MEI as the nearest 0.1 mg kets that he had ently, the Bau
rs searched Bol ulat ion is odu ctio n by these two snai ence of tem-
and uric acid (U), minus energy lost mined the ener as food. Since sites after heav linger’s study The eggs of l spec ies.
which is the nitro in feces (F) gy content of an aver he had deter- y rains. They each species
by lizards. We gen waste prod able to determin age cricket, Ang absent at a site concluded that temperatures—1 were incubate
can summarize uct produced e the energy inta illetta was only after two the snail was 98, 228, 258, and d at four
MEI in equation ing the number ke by each liza either a living
individual or an
2-hour surveys
failed to turn up pera ture s fall within the 29 8C. Not ice that these tem
form as: of crickets they rd by count- range measured -
MEI 5 C 2 F content of that ate and calculat The Baurs foun empty shell of (see fig. 5.34). by the satellite
2U number. He dete ing the energy the species. The eggs of both
Angilletta stud (F) and uric acid rmined the ener 29 sites surveyed d A. arbustorum still living 198C. However spec
, at higher tempera ies hatched at a high rate at
image
ied two populat (U) by collecti gy lost as feces by Bollinger near at 13 of the
South Carolina, ions from New produced by each ng all the fece remaining pop Basel. Eleven nificantly lowe tures, their eggs
regions with subs Jersey and lizard and then s and uric acid ulations lived in of these r rate s. At 228C, less hatc hed at sig-
tantially different material. He estim drying and wei two lived on gras deciduous fore eggs hatched, whi than 50% of A.
sy riverbanks. sts and le the eggs of C. arbustorum
climates.
and uric acid usin
ated the average
energy content
ghing this However, the Bau the other at a high rate. nemoralis cont
g a bomb calorime of feces rs could not At 258C, no A. inued to hatch
ter. approximately arbustorum eggs
50% of the C. hatched, while
nemoralis eggs
hatched. At 298
C,
moL37282_ch05_ moL37282_ch05_
099-124.indd 099-124.indd
106 122
7/23/14 5:54 7/23/14 5:55

PM PM
xvi Preface
New to the Seventh Edition rather than prey density per se. This discussion is coupled with
reviews of experimental and field studies that support the ratio-
The seventh edition expands the pedagogy by beginning dependent models.
all sections of every chapter with a list of student learn- The present edition connects ratio-dependent models
ing outcomes—over 450 student learning outcomes in all. of functional response to patterns of consumer abundance
These outcomes are largely based on fundamental learning and secondary production in ecosystems. Previous editions
outcomes for material covered in the text: have provided thorough coverage of the ecology of primary
1. Define key terms. production in terrestrial and aquatic ecosystems, but second-
2. Explain the main concepts. ary production has received much less attention. This seventh
3. Evaluate the strength of research presented in support of edition addresses this deficiency by including a section that
main concepts, including a critique of study design. covers the fundamentals of secondary production. The intro-
4. Interpret statistical evidence bearing on concepts, duction to secondary production in this edition is presented
expressed in graphical and numerical form. in the context of consumer responses to variations in primary
5. Apply the main concepts to interpretation of new production.
situations. New supplementary materials are placed online. Materi-
als cut from the sixth edition and those previously cut from the
A content thread focused on global change has been fifth and fourth editions are available online. Suggested read-
developed and distributed across chapters, emphasizing ings have been updated and placed online, along with answers to
global climate change. Students and instructors increasingly Concept Review and Critiquing the Evidence questions.
look for ways to connect the concepts and practice of ecologi-
cal science to environmental issues arising from global climate
Significant Chapter-by-Chapter Changes
change. The present edition explores how species are adjusting
their distributions and their critical life history events as cli- In chapters 1 to 23, numbered learning outcomes were
mate changes. The final chapter ends with a review of projected added to all concept discussions and Evaluating the Evidence
impacts of climate change on ecosystems and human popula- and Applications features. The average number of learning
tions, infrastructure, and economic systems. outcomes added to each chapter is 20.
This edition also builds on previous discussions of In chapter 10, a new Applications feature explores evi-
human disturbance of ecosystems to consider how damaged dence that plant and animal ranges have shifted northward and
ecosystems can be restored. The extent and intensity of human to higher latitudes in the Northern Hemisphere during the recent
impact on the biosphere grows with our population and expand- period of rapid global warming. This is the beginning of the
ing global economy. While climate change is the most promi- global climate change thread in the seventh edition. However, the
nent aspect of contemporary global change, other facets, such as presentation builds on earlier content in chapter 1 on population
damage or destruction of ecosystems, also call for solutions. As responses to climate change, including evolutionary responses,
a result, there is greater need to restore damaged communities and in chapter 4 on temperature relations of organisms.
and ecosystems. In this context, the new edition adds an intro- In chapter 12, a new Applications feature reviews studies
duction to the practice of ecological restoration, focusing on how that have shown shifts in the timing of flowering in plants and
the process of restoring ecosystems can benefit from concepts of migration in birds in response to climate warming. The dis-
developed in academic studies of community and ecosystem cussion complements the earlier discussion of shifts in species
succession. ranges in chapter 10 by demonstrating that climate warming is
The relationship between biodiversity and ecosystem not just inducing organisms to move in response to global warm-
function is introduced through the positive influence of pri- ing but also adjusting their life histories.
mary producer diversity on rates of primary production. In chapter 13, the Lotka-Volterra equations have been
Studies of biodiversity and ecosystem function are key elements modified from previous editions to make them more standard,
in ecology’s foundation. Connecting these elements helps create less cluttered, and easier for students to follow, which is essen-
conceptual coherence across the discipline. A growing body of tial, since these equations are the foundation of the mathematical
recent research does just that. Therefore, this edition includes a ecology covered in the text.
new section on the connection between biodiversity and ecosys- In chapter 14, we revisit predator functional responses
tem function. first introduced in chapter 7 by evaluating alternatives to those
The seventh edition introduces developments in trophic models. The Lotka-Volterra models of predator-prey interactions
ecology that build on classical models of predator-prey inter- published in the early twentieth century stimulated a long line
actions. The early to middle twentieth century was a golden of research. More recently, researchers have offered alternatives
age for theoretical ecology. However, those developments have that help identify where those classical mathematical models,
not stopped. Contemporary ecologists continue to build on that with their simplifying assumptions, apply and where alternative
legacy, improving our representation and understanding of eco- formulations better account for aspects of predator-prey inter-
logical systems as they do so. The seventh edition updates the actions, particularly at larger spatial and longer temporal
discussion of consumer functional response by introducing alter- scales. The discussion in this chapter reviews how recent ratio-
native models based on the ratio of prey to predator numbers dependent functional response models better predict predator
Preface xvii
functional responses in experimental and natural settings. The structure and function to these systems emerges as one of the
discussion helps to dispel the idea that mathematical ecology great contemporary ecological challenges. Increasingly ecolo-
ceased to develop in the mid-twentieth century and reinforces the gists addressing this challenge are turning to the conceptual
complementary roles of theoretical, experimental, and observa- framework of ecological succession to guide their work. Exam-
tional studies. ples of such work are included in this chapter to help bridge
In chapter 18, a new concept connects primary producer the historical divide between ecological theory and restoration
diversity to higher levels of primary production. The chapter also practice.
includes a new concept featuring the relationship between levels In chapter 23, the discussion of the Antarctic ozone hole
of primary production and secondary production. This discussion has been updated to 2013, including 35 years of data from NASA
provides a basis for introducing the fundamentals of secondary on the size of the ozone hole. The pattern shows that the maxi-
production. This addition also revisits the ratio-dependent func- mum size of the Antarctic ozone hole has stabilized, signaling
tional responses introduced in chapter 14 by extending the impli- a basis for ozone recovery predicted by atmospheric scientists
cations of those models beyond predator functional response to over the next 50 years, providing a bit of good planetary news.
the trophic structure of ecosystems. The treatment also formally The growing body of climate change research, published since
introduces secondary production, filling a conceptual gap in pre- the earlier editions of Ecology Concepts and Applications, has
vious editions. greatly improved understanding of how earth’s changing climate
In chapter 20, the fields of ecological restoration and will impact ecosystems and human populations, if not stabilized.
restoration ecology are introduced for the first time. Human A discussion of these impacts concludes this edition, underscor-
impact on the environment has altered ecological communities ing the relevance of ecological knowledge to sustaining natural
and ecosystems in nearly every corner of the planet. Restoring as well as human-centered systems.
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xix
xx Preface
Annual Editions: Environment 2015 Acknowledgments

by Eathorne
ISBN 978-1-25-916115-5 A complete list of the people who have helped me with this
Annual Editions is a compilation of current articles from the project would be impossibly long. However, during the devel-
best of the public press. The selections explore the global opment of this seventh edition, several colleagues freely
environment, the world’s energy, the biosphere, natural shared their ideas and expertise, reviewed new sections, or
resources, and pollution. Available through Create. offered the encouragement a project like this needs to keep
it going: Scott Collins, Cliff Dahm, Arturo Elosegi, Manuel
Graça, Tom Kennedy, Tim Lowrey, Sam Loker, Rob Miller,
Taking Sides: Clashing Views on Will Pockman, Steve Poe, Bob Sinsabaugh, Alain Thomas,
Environmental Issues, Tom Turner, Lawrence Walker, Chris Witt, Blair Wolf. I wish
Sixteenth Edition by Easton to offer special thanks to Roger Arditi and Lev Ginzburg
ISBN: 978-1-25-916113-1 for their time and patience in helping me develop sections
Taking Sides presents current contro- on ratio-dependent models of functional response and their
versial issues in a debate-style format potential contributions to better understanding of predator-
designed to stimulate student interest prey interactions and the trophic structure of ecosystems. I am
and develop critical thinking skills. also grateful to Art Benke for helping me develop an over-
Each issue is thoughtfully framed with view of secondary production for this edition and for helping
an issue summary, an issue introduction, and a postscript or integrate it with discussion of the effects of enrichment on
challenge questions. An online Instructor’s Resource Guide ecosystem trophic structure. John and Leah Vucetich helped
with testing material is available. Available through Create. bring their long-term research on wolf-moose interactions on
Isle Royale to life by graciously allowing use of one of their
Classic Edition Sources: Environmental Studies many photos of interactions in this model predator and prey
Fourth Edition by Thomas Easton system. In addition, I am indebted to the many students and
ISBN 978-0-07-352764-2 instructors who have helped by contacting me with questions
Sources brings together selections of enduring intellectual and suggestions for improvements.
value—classic articles, book excerpts, and research studies— I also wish to acknowledge the skillful guidance and work
that have shaped ecology and environmental science. Edited throughout the publishing process given by many profession-
for length and level, the selections are organized topically. als associated with McGraw-Hill during this project, including
An annotated table of contents provides a quick and easy Becky Olson, Patrick Reidy, Carrie Burger, Fran Simon, April
review of the selections. Supported by an online instructor’s Southwood, Lynn Breithaupt, Mary Reeg, Angie Sigwarth, Tara
Resource Guide that provides a complete synopsis of each selec- McDermott, and Sheila Frank.
tion, guidelines for discussing the selection in class, and testing Finally, I wish to thank all my family for support given
materials. Available through Create. throughout the project, especially Paulette Dompeling, Mary Ann
Esparza, Dan Esparza, Hani Molles, Anders Molles, Mary Anne
Ecology Laboratory Manual, by Vodopich Nelson, and Keena.
(ISBN: 978-0-07-338318-7; I gratefully acknowledge the many reviewers who, over the
MHID: 0-07-338318-X) course of the last several revisions, have given of their time and
Darrell Vodopich, co-author of Biology Laboratory Manual, expertise to help this textbook evolve to its present seventh edi-
has written a new lab manual for ecology. This lab manual tion. Their depth and breadth of knowledge and experience, both
offers straightforward procedures that are doable in a broad as researchers and teachers, are humbling. They continue my
range of classroom, lab, and field situations. The procedures education, for which I am grateful, and I honestly could not have
have specific instructions that can be taught by a teaching continued the improvement of this textbook without them.
assistant with minimal experience as well as by a professor. I gratefully acknowledge the many reviewers who, over
the course of the last several revisions, have given of their
Student Atlas of Environmental time and expertise to help this textbook evolve to its present
Issues, by Allen edition. Their depth and breadth of knowledge and experi-
(ISBN: 978-0-69-736520-0; ence, both as researchers and teachers, are humbling. They
MHID: 0-69-736520-4) continue my education, for which I am grateful, and I hon-
This atlas is an invaluable pedagogical estly could not have continued the improvement of this text-
tool for exploring the human impact on book without them.
the air, waters, biosphere, and land in
every major world region. This infor- Reviewers for the Seventh Edition
mative resource provides a unique John Bacheller Hillsborough Community College
combination of maps and data that help students understand Isaac Barjis City University of New York
the dimensions of the world’s environmental problems and Dena Berg Tarrant County College NW
the geographic basis of these problems. Earl R. Beyer Harrisburg Area Community College
Preface xxi
Jamal Bittar The University of Toledo Jerry Baskin University of Kentucky

Linda Bruslind Oregon State University Thomas O. Crist Miami University
Sherri L. Buerdsell West Virginia Northern Community College Peter Alpert University of Massachusetts—Amherst
Carrie E. Burdzinski Delta College (University Center, Michigan) Mark Pyron Ball State University
William Dew Nipissing University Mary Bremigan Michigan State University
Harry G. Deneer University of Saskatchewan
Phil Denette Delgado Community College Reviewers for the Fifth Edition
Jessica A. DiGirolamo Broward College, Davie, Florida
Angela M. Edwards Trident Technical College Joel S. Brown University of Illinois—Chicago
Elyce Ervin University of Toledo Peter E. Busher Boston University
Teresa G. Fischer Indian River State College Lloyd Fitzpatrick University of North Texas
Christina Gan Highline Community College James A. Fordyce University of Tennessee
Kathryn Germain Southwest Tennessee Community College David L. Gorchov Miami University
Linda Girouard Brescia University Jamie Kneitel California State University—Sacramento
Judy Gnarpe University of Alberta John C. Krenetsky Metropolitan State College of Denver
Amy D. Goode Illinois Central College Amy E. Lesen Pratt Institute
Robert C. Hairston Harrisburg Area Community College D. Nicholas McLetchie University of Kentucky
Nasreen S. Haque City University of New York, New York Thomas Pliske Florida International University
Daniel P. Herman University of Wisconsin—Eau Claire Nathan J. Sanders University of Tennessee
Ingrid Herrmann Santa Fe College Robert M. Schoch Boston University
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Chike Igboechi Medgar Evers College of the City University
of New York Reviewers for the Fourth Edition
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xxii Preface
Craig E. Williamson Miami University of Ohio Thomas W. Jurik Iowa State University
Jianguo (Jingle) Wu Arizona State University Karen L. Kandl University of New Orleans
Douglas Zook Boston University Robert Keys Cornerstone University
Mark E. Knauss Shorter College
Reviewers for the Third Edition Jean Knops University of Nebraska
Anthony J. Krzysik Embry-Riddle Aeronautical University
Sina Adl Dalhousie University, Canada Eddie N. Laboy-Nieves InterAmerican University
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Peter Alpert University of Massachusetts—Amherst Vic Landrum Washburn University
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Edmund Bedecarrax City College of San Francisco John S. Mackiewicz State University of New York—Albany
Jerry Beilby Northwestern College Tim Maret Shippensburg University
R. P. Benard American International College Ken R. Marion University of Alabama—Birmingham
Erica Bergquist Holyoke Community College Vicky Meretsky Indiana University
Richard A. Boutwell Missouri Western State College John C. Mertz Delaware Valley College
Ward Brady Arizona State University East—Mesa Carolyn Meyer University of Wyoming
Fred J. Brenner Grove City College Sheila G. Miracle Southeast Community College—Bell City
Robert Brodman Saint Joseph’s College Timothy Mousseau University of South Carolina
Elaine R. Brooks San Diego City College Virginia Naples Northern Illinois University
Evert Brown Casper College Peter Nonacs University of California—Los Angeles
Stephanie Brown Fabritius Southwestern University Mark H. Olson Franklin and Marshall College
Rebecca S. Burton Alverno College David W. Onstad University of Illinois—Champaign
James E. Byers University of New Hampshire Fatimata A. Palé Thiel College
Guy Cameron University of Cincinnati Mary Lou Peltier Saint Martin’s College
Geralyn M. Caplan Owensboro Community Carolyn Peters Spoon River College
and Technical College Kenneth L. Petersen Dordt College
Walter P. Carson University of Pittsburgh Eric R. Pianka University of Texas
Ben Cash III Maryville College Raymond Pierotti University of Kansas—Lawrence
Young D. Choi Purdue University—Calumet David Pindel Corning Community College
Ethan Clotfelter Providence College Jon K. Piper Bethel College
Liane Cochran-Stafira Saint Xavier University Thomas E. Pliske Florida International University
Joe Coelho Culver-Stockton College Michael V. Plummer Harding University
Jerry L. Cook Sam Houston State University Ellen Porter Holtman Virginia Western Community College
Tamara J. Cook Sam Houston State University Diane Post University of Texas—Permian Basin
Erica Corbett Southeastern Oklahoma State University Kathleen Rath Marr Lakeland College
Tim Craig University of Minnesota Brian C. Reeder Morehead State University
Jack A. Cranford Virginia Tech Seth R. Reice University of North Carolina—Chapel Hill
Greg Cronin University of Colorado—Denver Robin Richardson Winona State University
Todd Crowl Utah State University Carol D. Riley Gainesville College
Richard J. Deslippe Texas Tech University Marianne W. Robertson Millikin University
Kenneth M. Duke Brevard College Tom Robertson Portland Community College
Andy Dyer University of South Carolina Bernadette M. Roche Loyola College in Maryland
Ginny L. Eckert University of Alaska Tatiana Roth Coppin State College
J. Nicholas Ehringer Hillsborough Community College Neil Sabine Indiana University East
George F. Estabrook University of Michigan Seema Sanjay Jejurikar Bellevue Community College
Richard S. Feldman Marist College Timothy Savisky University of Pittsburgh
Charles A. Francis University of Nebraska—Lincoln Josh Schimel University of California—Santa Barbara
Carl Freeman Wayne State University Michael G. Scott Lincoln University
J. Phil Gibson Agnes Scott College Erik R. Scully Towson University
Robert R. Glesener Brevard College Michael J. Sebetich William Paterson University
Michael L. Golden Grossmont College Walter M. Shriner Mount Hood Community College
Paul Grecay Salisbury University John Skillman California State University—San Bernardino
Lana Hamilton Northeast State Tech Community College Jerry M. Skinner Keystone College
Brian Helmuth University of South Carolina Garriet W. Smith University of South Carolina—Aiken
James R. Hodgson Saint Norbert College Stacy Smith Lexington Community College
Jeremiah N. Jarrett Central Connecticut State University Joseph Stabile Iona College
Krish Jayachandran Florida International University Alan Stam Capital University
Mark Jonasson Crafton Hills College Alan Stiven University of North Carolina—Chapel Hill
Preface xxiii
Eric D. Storie Roanoke-Chowan Community College Fred E. Wasserman Boston University

William A. Szelistowski Eckerd College Phillip L. Watson Ferris State University
Robert Tatina Dakota Wesleyan University Donna Wear Augusta State University
Nina N. Thumser California University of Pennsylvania John F. Wegner Emory State University
John A. Tiedemann Monmouth University Matt R. Whiles Southern Illinois University
Anne H. Todd Bockarie Philadelphia University Howard Whiteman Murray State University
Conrad Toepfer Millikin University Craig E. Williamson Lehigh University
Donald E. Trisel Fairmont State College Gordon Wolfe California State University—Chico
Dessie L. A. Underwood California State University— Derek Zelmer Emporia State University
Long Beach Douglas Zook Boston University
Carl Von Ende Northern Illinois University Manuel C. Molles Jr.
Introduction
to Ecology
1
Historical Foundations
and Developing Frontiers
A yellow-rumped warbler, Dendroica coronata, feeding young.
Ecological studies of warblers have made fundamental contributions LEARNING OUTCOME
to the growth of ecological understanding. After studying this section you should be able to do the following:
1.1 Discuss the concept of environment as it pertains to

the science of ecology.
W
hat is ecology? Ecology, the study of relation-
CHAPTER CONCEPTS ships between organisms and the environment,
has been a focus for human study for as long as
1.1 Ecologists study environmental we have existed as a species. Our survival has depended upon
relationships ranging from those how well we could observe variations in the environment and
of individual organisms to factors predict the responses of organisms to those variations. The
influencing global-scale processes. 2 earliest hunters and gatherers had to know the habits of their
Concept 1.1 Review 3 animal prey and where to find food plants. Later, agricultur-
ists had to be aware of variations in weather and soils and of
1.2 Ecologists design their studies based on how such variation might affect crops and livestock.
their research questions, the temporal Today, most of earth’s human population live in cities and
and spatial scale of their studies, and most of us have little direct contact with nature. More than ever
available research tools. 3 before, though, the future of our species depends on how well we
Concept 1.2 Review 8 understand the relationships between organisms and the environ-
ment. Our species is rapidly changing earth’s environment, yet
Investigating the Evidence 1: we do not fully understand the consequences of these changes.
The Scientific Method—Questions For instance, human activity has increased the quantity of nitro-
and Hypotheses 9 gen cycling through the biosphere, changed land cover across
Summary 10 the globe, and increased the atmospheric concentration of CO2.
Key Terms 10 Changes such as these threaten the diversity of life on earth and
Review Questions 10 may endanger our life support system. Because of the rapid pace
of environmental change at the dawn of the twenty-first century,
it is imperative that we continue as ardent students of ecology.
1
Another random document with
no related content on Scribd:
It may be replied that the history of Rome, on which I dwelt a
moment ago, shews that arrested progress, and even decadence,
may be but the prelude to a new period of vigorous growth. So that
even those races or nations which seem frozen into eternal
immobility may base upon experience their hopes of an awakening
spring.
I am not sure, however, that this is the true interpretation of the facts.
There is no spectacle indeed in all history more impressive than the
thick darkness settling down over Western Europe, blotting out all
but a faint and distorted vision of Graeco-Roman culture, and then,
as it slowly rises, unveiling the variety and rich promise of the
modern world. But I do not think we should make this unique
phenomenon support too weighty a load of theory. I should not infer
from it that when some wave of civilisation has apparently spent its
force, we have a right to regard its withdrawing sweep as but the
prelude to a new advance. I should rather conjecture that in this
particular case we should find, among other subtle causes of
decadence, some obscure disharmony between the Imperial system
and the temperament of the West, undetected even by those who
suffered from it. That system, though accepted with contentment and
even with pride, though in the days of its greatness it brought
civilisation, commerce, and security in its train, must surely have
lacked some elements which are needed to foster among Teutons,
Celts, and Iberians the qualities, whatever these may be, on which
sustained progress depends. It was perhaps too oriental for the
occident, and it certainly became more oriental as time went on. In
the East it was, comparatively speaking, successful. If there was no
progress, decadence was slow; and but for what Western Europe
did, and what it failed to do, during the long struggle with militant
Mahommedanism, there might still be an Empire in the East, largely
Asiatic in population, Christian in religion, Greek in culture, Roman
by political descent.
Had this been the course of events large portions of mankind would
doubtless have been much better governed than they are. It is not so
clear that they would have been more ‘progressive.’ Progress is with
the West: with communities of the European type. And if their energy
of development is some day to be exhausted, who can believe that
there remains any external source from which it can be renewed?
Where are the untried races competent to construct out of the ruined
fragments of our civilisation a new and better habitation for the spirit
of man? They do not exist: and if the world is again to be buried
under a barbaric flood, it will not be like that which fertilised, though it
first destroyed, the western provinces of Rome, but like that which in
Asia submerged for ever the last traces of Hellenic culture.
We are thus brought back to the question I put a few moments since.
What grounds are there for supposing that we can escape the fate to
which other races have had to submit? If for periods which,
measured on the historic scale, are of great duration, communities
which have advanced to a certain point appear able to advance no
further; if civilisations wear out, and races become effete, why should
we expect to progress indefinitely, why for us alone is the doom of
man to be reversed?
To these questions I have no very satisfactory answers to give, nor
do I believe that our knowledge of national or social psychology is
sufficient to make a satisfactory answer possible. Some purely
tentative observations on the point may, however, furnish a fitting
conclusion to an address which has been tentative throughout, and
aims rather at suggesting trains of thought, than at completing them.
I assume that the factors which combine to make each generation
what it is at the moment of its entrance into adult life are in the main
twofold. The one produces the raw material of society, the process of
manufacture is effected by the other. The first is physiological
inheritance, the second is the inheritance partly of external
conditions of life, partly of beliefs[2], traditions, sentiments, customs,
laws, and organisation—all that constitute the social surroundings in
which men grow up to maturity.
I hazard no conjecture as to the share borne respectively by these
two kinds of cause in producing their joint result. Nor are we likely to
obtain satisfactory evidence on the subject till, in the interests of
science, two communities of different blood and different traditions
consent to exchange their children at birth by a universal process of
reciprocal adoption. But even in the absence of so heroic an
experiment, it seems safe to say that the mobility which makes
possible either progress or decadence, resides rather in the causes
grouped under the second head than in the physiological material on
which education, in the widest sense of that ambiguous term, has
got to work. If, as I suppose, acquired qualities are not inherited, the
only causes which could fundamentally modify the physiological
character of any particular community are its intermixture with alien
races through slavery, conquest, or immigration; or else new
conditions which varied the relative proportion in which different
sections of the population contributed to its total numbers. If, for
example, the more successful members of the community had
smaller families than the less successful; or if medical administration
succeeded in extinguishing maladies to which persons of a particular
constitution were specially liable; or if one strain in a mixed race had
a larger birth rate than another—in these cases and in others like
them, there would doubtless be a change in the physiological factor
of national character. But such changes are not likely, I suppose, to
be considerable, except, perhaps, those due to the mixture of races;
—and that only in new countries whose economic opportunities
tempt immigrants widely differing in culture, and in capacity for
culture, from those whose citizenship they propose to share.
The flexible element in any society, that which is susceptible of
progress or decadence, must therefore be looked for rather in the
physical and psychical conditions affecting the life of its component
units, than in their inherited constitution. This last rather supplies a
limit to variations than an element which does itself vary: though
from this point of view its importance is capital. I at least find it quite
impossible to believe that any attempt to provide widely different
races with an identical environment, political, religious, educational,
what you will, can ever make them alike. They have been different
and unequal since history began; different and unequal they are
destined to remain through future periods of comparable duration.
But though the advance of each community is thus limited by its
inherited aptitudes, I do not suppose that those limits have ever been
reached by its unaided efforts. In the cases where a forward
movement has died away, the pause must in part be due to arrested
development in the variable, not to a fixed resistance in the
unchanging factor of national character. Either external conditions
are unfavourable; or the sentiments, customs and beliefs which
make society possible have hardened into shapes which make its
further self-development impossible; or through mere weariness of
spirit the community resigns itself to a contented, or perhaps a
discontented, stagnation; or it shatters itself in pursuit of impossible
ideals, or for other and obscurer reasons, flags in its endeavours,
and falls short of possible achievement.
Now I am quite unable to offer any such general analysis of the
causes by which these hindrances to progress are produced or
removed as would furnish a reply to my question. But it may be
worth noting that a social force has come into being, new in
magnitude if not in kind, which must favourably modify such
hindrances as come under all but the last of the divisions in which I
have roughly arranged them. This force is the modern alliance
between pure science and industry. That on this we must mainly rely
for the improvement of the material conditions under which societies
live is in my opinion obvious, although no one would conjecture it
from a historic survey of political controversy. Its direct moral effects
are less obvious; indeed there are many most excellent people who
would altogether deny their existence. To regard it as a force fitted to
rouse and sustain the energies of nations would seem to them
absurd: for this would be to rank it with those other forces which
have most deeply stirred the emotions of great communities, have
urged them to the greatest exertions, have released them most
effectually from the benumbing fetters of merely personal
preoccupations,—with religion, patriotism, and politics. Industrial
expansion under scientific inspiration, so far from deserving praise
like this, is in their view, at best, but a new source of material well-
being, at worst the prolific parent of physical ugliness in many forms,
machine made wares, smoky cities, polluted rivers, and desecrated
landscapes,—appropriately associated with materialism and greed.
I believe this view to be utterly misleading, confounding accident with
essence, transient accompaniments with inseparable characteristics.
Should we dream of thus judging the other great social forces of
which I have spoken? Are we to ignore what religion has done for the
world because it has been the fruitful excuse for the narrowest
bigotries and the most cruel persecutions? Are we to underrate the
worth of politics, because politics may mean no more than the
mindless clash of factions, or the barren exchange of one set of
tyrants or jobbers for another? Is patriotism to be despised because
its manifestations have been sometimes vulgar, sometimes selfish,
sometimes brutal, sometimes criminal? Estimates like these seem to
me worse than useless. All great social forces are not merely
capable of perversion, they are constantly perverted. Yet were they
eliminated from our social system, were each man, acting on the
advice, which Voltaire gave but never followed, to disinterest himself
of all that goes on beyond the limits of his own cabbage garden,
decadence I take it, would have already far advanced.
But if the proposition I am defending may be wrongly criticised, it is
still more likely to be wrongly praised. To some it will commend itself
as a eulogy on an industrial as distinguished from a military
civilisation: as a suggestion that in the peaceful pursuit of wealth
there is that which of itself may constitute a valuable social tonic.
This may be true, but it is not my contention. In talking of the alliance
between industry and science my emphasis is at least as much on
the word science as on the word industry. I am not concerned now
with the proportion of the population devoted to productive labour, or
the esteem in which they are held. It is on the effects which I believe
are following, and are going in yet larger measure to follow, from the
intimate relation between scientific discovery and industrial
efficiency, that I most desire to insist.
Do you then, it will be asked, so highly rate the utilitarian aspect of
research as to regard it as a source, not merely of material
convenience, but of spiritual elevation? Is it seriously to be ranked
with religion and patriotism as an important force for raising men’s
lives above what is small, personal, and self-centred? Does it not
rather pervert pure knowledge into a new contrivance for making
money, and give a fresh triumph to the ‘growing materialism of the
age’?
I do not myself believe that this age is either less spiritual or more
sordid than its predecessors. I believe, indeed, precisely the reverse.
But however this may be, is it not plain that if a society is to be
moved by the remote speculations of isolated thinkers it can only be
on condition that their isolation is not complete? Some point of
contact they must have with the world in which they live, and if their
influence is to be based on widespread sympathy, the contact must
be in a region where there can be, if not full mutual comprehension,
at least a large measure of practical agreement and willing co-
operation. Philosophy has never touched the mass of men except
through religion. And, though the parallel is not complete, it is safe to
say that science will never touch them unaided by its practical
applications. Its wonders may be catalogued for purposes of
education, they may be illustrated by arresting experiments, by
numbers and magnitudes which startle or fatigue the imagination;
but they will form no familiar portion of the intellectual furniture of
ordinary men unless they be connected, however remotely, with the
conduct of ordinary life. Critics have made merry over the naive self-
importance which represented man as the centre and final cause of
the universe, and conceived the stupendous mechanism of nature as
primarily designed to satisfy his wants and minister to his
entertainment. But there is another, and an opposite, danger into
which it is possible to fall. The material world, howsoever it may have
gained in sublimity, has, under the touch of science, lost (so to
speak) in domestic charm. Except where it affects the immediate
needs of organic life, it may seem so remote from the concerns of
men that in the majority it will rouse no curiosity, while of those who
are fascinated by its marvels, not a few will be chilled by its
impersonal and indifferent immensity.
For this latter mood only religion or religious philosophy can supply a
cure. But for the former, the appropriate remedy is the perpetual
stimulus which the influence of science on the business of mankind
offers to their sluggish curiosity. And even now I believe this
influence to be underrated. If in the last hundred years the whole
material setting of civilised life has altered, we owe it neither to
politicians nor to political institutions. We owe it to the combined
efforts of those who have advanced science and those who have
applied it. If our outlook upon the Universe has suffered
modifications in detail so great and so numerous that they amount
collectively to a revolution, it is to men of science we owe it, not to
theologians or philosophers. On these indeed new and weighty
responsibilities are being cast. They have to harmonise and to
coordinate, to prevent the new from being one-sided, to preserve the
valuable essence of what is old. But science is the great instrument
of social change, all the greater because its object is not change but
knowledge; and its silent appropriation of this dominant function,
amid the din of political and religious strife, is the most vital of all the
revolutions which have marked the development of modern
civilisation.
It may seem fanciful to find in a single recent aspect of this revolution
an influence which resembles religion or patriotism in its appeals to
the higher side of ordinary characters—especially since we are
accustomed to regard the appropriation by industry of scientific
discoveries merely as a means of multiplying the material
conveniences of life. But if it be remembered that this process brings
vast sections of every industrial community into admiring relation
with the highest intellectual achievement, and the most disinterested
search for truth; that those who live by ministering to the common
wants of average humanity lean for support on those who search
among the deepest mysteries of Nature; that their dependence is
rewarded by growing success; that success gives in its turn an
incentive to individual effort in no wise to be measured by personal
expectation of gain; that the energies thus aroused may affect the
whole character of the community, spreading the beneficent
contagion of hope and high endeavour through channels scarcely
known, to workers[3] in fields the most remote; if all this be borne in
mind it may perhaps seem not unworthy of the place I have assigned
to it.
But I do not offer this speculation, whatever be its worth, as an
answer to my original question. It is but an aid to optimism, not a
reply to pessimism. Such a reply can only be given by a sociology
which has arrived at scientific conclusions on the life-history of
different types of society, and has risen above the empirical and
merely interrogative point of view which, for want of a better, I have
adopted in this address. No such sociology exists at present, or
seems likely soon to be created. In its absence the conclusions at
which I provisionally arrive are that we cannot regard decadence and
arrested development as less normal in human communities than
progress; though the point at which the energy of advance is
exhausted (if, and when it is reached) varies in different races and
civilisations: that the internal causes by which progress is
encouraged, hindered, or reversed, lie to a great extent beyond the
field of ordinary political discussion, and are not easily expressed in
current political terminology: that the influence which a superior
civilisation, whether acting by example or imposed by force, may
have in advancing an inferior one, though often beneficent, is not
likely to be self supporting; its withdrawal will be followed by
decadence, unless the character of the civilisation be in harmony
both with the acquired temperament and the innate capacities of
those who have been induced to accept it: that as regards those
nations which still advance in virtue of their own inherent energies,
though time has brought perhaps new causes of disquiet, it has
brought also new grounds of hope; and that whatever be the perils in
front of us, there are, so far, no symptoms either of pause or of
regression in the onward movement which for more than a thousand
years has been characteristic of Western civilisation.
Notes:
[1] The ‘East’ is a term most loosely used. It does not here include
China and Japan and does include parts of Africa. The observations
which follow have no reference either to the Jews or to the
commercial aristocracies of Phœnician origin.
[2] Beliefs include knowledge.
[3] This remark arises out of a train of thought suggested by two
questions which are very pertinent to the subject of the Address.
(1) Is a due succession of men above the average in original
capacity necessary to maintain social progress? and
(2) If so, can we discover any law according to which such men are
produced?
I entertain no doubt myself that the answer to the first question
should be in the affirmative. Democracy is an excellent thing; but,
though quite consistent with progress, it is not progressive per se. Its
value is regulative not dynamic; and if it meant (as it never does)
substantial uniformity, instead of legal equality, we should become
fossilised at once. Movement may be controlled or checked by the
many; it is initiated and made effective by the few. If (for the sake of
illustration) we suppose mental capacity in all its many forms to be
mensurable and commensurable, and then imagine two societies
possessing the same average capacity—but an average made up in
one case of equal units, in the other of a majority slightly below the
average and a minority much above it, few could doubt that the
second, not the first, would show the greatest aptitude for
movement. It might go wrong, but it would go.
The second question—how is this originality (in its higher
manifestations called genius) effectively produced? is not so simple.
Excluding education in its narrowest sense—which few would regard
as having much to do with the matter—the only alternatives seem to
be the following:
Original capacity may be no more than one of the ordinary variations
incidental to heredity. A community may breed a minority thus
exceptionally gifted, as it breeds a minority of men over six feet six.
There may be an average decennial output of congenital geniuses
as there is an average decennial output of congenital idiots—though
the number is likely to be smaller.
But if this be the sole cause of the phenomenon, why does the same
race apparently produce many men of genius in one generation and
few in another? Why are years of abundance so often followed by
long periods of sterility?
The most obvious explanation of this would seem to be that in some
periods circumstances give many openings to genius, in some
periods few. The genius is constantly produced; but it is only
occasionally recognised.
In this there must be some truth. A mob orator in Turkey, a religious
reformer in seventeenth century Spain, a military leader in the
Sandwich islands, would hardly get their chance. Yet the theory of
opportunity can scarcely be reckoned a complete explanation. For it
leaves unaccounted for the variety of genius which has in some
countries marked epochs of vigorous national development. Athens
in the fifth and fourth centuries, Florence in the fifteenth and early
sixteenth centuries, Holland in the later sixteenth and seventeenth
centuries, are the typical examples. In such periods the opportunities
of statesmen, soldiers, orators, and diplomatists, may have been
specially frequent. But whence came the poets, the sculptors, the
painters, the philosophers and the men of letters? What peculiar
opportunities had they?
The only explanation, if we reject the idea of a mere coincidence,
seems to be, that quite apart from opportunity, the exceptional stir
and fervour of national life evokes or may evoke qualities which in
ordinary times lie dormant, unknown even to their possessors. The
potential Miltons are ‘mute’ and ‘inglorious’ not because they cannot
find a publisher, but because they have nothing they want to publish.
They lack the kind of inspiration which, on this view, flows from social
surroundings where great things, though of quite another kind, are
being done and thought.
If this theory be true (and it is not without its difficulties) one would
like to know whether these undoubted outbursts of originality in the
higher and rarer form of genius, are symptomatic of a general rise in
the number of persons exhibiting original capacity of a more ordinary
type. If so, then the conclusion would seem to be that some kind of
widespread exhilaration or excitement is required in order to enable
any community to extract the best results from the raw material
transmitted to it by natural inheritance.
Cambridge: Printed at the University Press.
Transcriber’s Note
The formatting of the notes was substantially altered for this edition.
On page 41, “Greek in culture Roman by political descent” was corrected to “Greek in
culture, Roman by political descent.”
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8.2 Mate Choice and Resource Provisioning

8.4 Sociality 184

Concepts 258 Applications: Competition between Native

Section IV Models 317

15.3 Evolution of Mutualism 344 17.2 Indirect Interactions 376

Chapter 20 Succession and Stability 435

Concepts 435 Structure 472

Manipulating Island Area 493 El Niño and Marine Populations 511

ecology and includes chapters on landscape, geographic,

section I within a natural history context. In summary, the

historical events related to the subject; others pre- moL37282_ch05_

sent an example of an ecological process. All attempt 99

students with key information where they Birds leave the

of on the y-axis creates a tio

pi- Dall sheep surviving their

100 wolves and die

Confirming Pag Confirming Pag

7/23/14 5:54 7/23/14 5:55

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Annual Editions: Environment 2015 Acknowledgments

Jamal Bittar The University of Toledo Jerry Baskin University of Kentucky

Eric D. Storie Roanoke-Chowan Community College Fred E. Wasserman Boston University

1.1 Discuss the concept of environment as it pertains to

Cambridge: Printed at the University Press.

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