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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

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

Chapter 12 Life Histories 258 Investigating the Evidence 13: Field Experiments
Concept 13.4 Review 300
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:
Concept 14.2 Review 312
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

Concept 14.3 Review 319
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
Concept 13.1 Review 286 Concept 14.5 Review 326
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
Concept 13.2 Review 289
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
Concept 15.3 Review 348 Concept 17.2 Review 378
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
Concept 17.3 Review 386
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
Concept 16.2 Review 357
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
Concept 16.3 Review 364
Concept 18.2 Review 398
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
Concept 18.3 Review 400
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
Concept 17.1 Review 375 Concept 18.5 Review 409
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
Concept 20.4 Review 453
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
Concept 19.1 Review 419
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
Concept 19.2 Review 424
19.3 Organisms and Nutrients 425 21.1 Landscape Structure 462

Nutrient Cycling in Streams and Lakes 425
Animals and Nutrient Cycling in Terrestrial
Ecosystems 427
Plants and the Nutrient Dynamics of Ecosystems
Concept 19.3 Review 429
19.4 Disturbance and Nutrients 429
Disturbance and Nutrient Loss from Forests 429
Flooding and Nutrient Export by Streams 430
Concept 19.4 Review 431
Applications: Altering Aquatic and Terrestrial
Ecosystems 432
The Structure of Six Landscapes in Ohio 462
The Fractal Geometry of Landscapes 464
Concept 21.1 Review 465
428 21.2 Landscape Processes 465
Landscape Structure and the Dispersal of Mammals 466
Habitat Patch Size and Isolation and the Density
of Butterfly Populations 467
Habitat Corridors and Movement of Organisms 468
Landscape Position and Lake Chemistry 469
Investigating the Evidence 21: Comparison of Two Samples
Using a Rank Sum Test 470
Concept 21.2 Review 471

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
Concept 20.1 Review 440
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
Concept 20.2 Review 446 Concept 22.1 Review 489
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

This book was written for students taking their first under-
graduate course in ecology. I have assumed that students
in this one-semester course have some knowledge of basic
chemistry and mathematics and have had a course in general
biology, which included introductions to physiology, biologi-
cal diversity, and evolution.
Organization of the Book
An evolutionary perspective forms the foundation of the
entire textbook, as it is needed to support understanding
of major concepts. The textbook begins with a brief intro-
duction to the nature and history of the discipline of ecol-
ogy, followed by section I, which includes two chapters on
natural history—life on land and life in water and a chapter
on population genetics and natural selection. Sections II
through VI build a hierarchical perspective through the
traditional subdisciplines of ecology: section II concerns
adaptations to the environment; section III focuses on
population ecology; section IV presents the ecology of
interactions; section V summarizes community and ecosys-
tem ecology; and finally, section VI discusses large-scale
Preface
to engage students and draw them into the discussion that
follows.
Concepts: The goal of this book is to build a foundation of
ecological knowledge around key concepts. I have found that
while beginning ecology students can absorb a few central
concepts well, they can easily get lost in a sea of details. The
key concepts are listed at the beginning of each chapter to
alert the student to the major topics to follow and to provide a
place where the student can find a list of the important points
covered in each chapter. The sections in which concepts are
discussed focus on published studies and, wherever possible,
the scientists who did the research are introduced. This case-
study approach supports the concepts with evidence, and
introduces students to the methods and people that have cre-
ated the discipline of ecology. Each concept discussion ends
with a series of concept review questions to help students
test their knowledge and to reinforce key points made in the
discussion.
Confirming Pag
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
The seventh edition expands the pedagogy by beginning
all sections of every chapter with a list of student learn-
ing outcomes—over 450 student learning outcomes in all.
These outcomes are largely based on fundamental learning
outcomes for material covered in the text:
1. Define key terms.
2. Explain the main concepts.
3. Evaluate the strength of research presented in support of
main concepts, including a critique of study design.
4. Interpret statistical evidence bearing on concepts,
expressed in graphical and numerical form.
5. Apply the main concepts to interpretation of new
situations.
A content thread focused on global change has been
developed and distributed across chapters, emphasizing
global climate change. Students and instructors increasingly
look for ways to connect the concepts and practice of ecologi-
cal science to environmental issues arising from global climate
change. The present edition explores how species are adjusting
their distributions and their critical life history events as cli-
mate changes. The final chapter ends with a review of projected
impacts of climate change on ecosystems and human popula-
tions, infrastructure, and economic systems.
This edition also builds on previous discussions of
human disturbance of ecosystems to consider how damaged
ecosystems can be restored. The extent and intensity of human
impact on the biosphere grows with our population and expand-
ing global economy. While climate change is the most promi-
nent aspect of contemporary global change, other facets, such as
damage or destruction of ecosystems, also call for solutions. As
a result, there is greater need to restore damaged communities
and ecosystems. In this context, the new edition adds an intro-
duction to the practice of ecological restoration, focusing on how
the process of restoring ecosystems can benefit from concepts
developed in academic studies of community and ecosystem
succession.
The relationship between biodiversity and ecosystem
function is introduced through the positive influence of pri-
mary producer diversity on rates of primary production.
Studies of biodiversity and ecosystem function are key elements
in ecology’s foundation. Connecting these elements helps create
conceptual coherence across the discipline. A growing body of
recent research does just that. Therefore, this edition includes a
new section on the connection between biodiversity and ecosys-
tem function.
The seventh edition introduces developments in trophic
ecology that build on classical models of predator-prey inter-
actions. The early to middle twentieth century was a golden
age for theoretical ecology. However, those developments have
not stopped. Contemporary ecologists continue to build on that
legacy, improving our representation and understanding of eco-
logical systems as they do so. The seventh edition updates the
discussion of consumer functional response by introducing alter-
native models based on the ratio of prey to predator numbers
rather than prey density per se. This discussion is coupled with
reviews of experimental and field studies that support the ratio-
dependent models.
The present edition connects ratio-dependent models
of functional response to patterns of consumer abundance
and secondary production in ecosystems. Previous editions
have provided thorough coverage of the ecology of primary
production in terrestrial and aquatic ecosystems, but second-
ary production has received much less attention. This seventh
edition addresses this deficiency by including a section that
covers the fundamentals of secondary production. The intro-
duction to secondary production in this edition is presented
in the context of consumer responses to variations in primary
production.
New supplementary materials are placed online. Materi-
als cut from the sixth edition and those previously cut from the
fifth and fourth editions are available online. Suggested read-
ings have been updated and placed online, along with answers to
Concept Review and Critiquing the Evidence questions.
Significant Chapter-by-Chapter Changes
In chapters 1 to 23, numbered learning outcomes were
added to all concept discussions and Evaluating the Evidence
and Applications features. The average number of learning
outcomes added to each chapter is 20.
In chapter 10, a new Applications feature explores evi-
dence that plant and animal ranges have shifted northward and
to higher latitudes in the Northern Hemisphere during the recent
period of rapid global warming. This is the beginning of the
global climate change thread in the seventh edition. However, the
presentation builds on earlier content in chapter 1 on population
responses to climate change, including evolutionary responses,
and in chapter 4 on temperature relations of organisms.
In chapter 12, a new Applications feature reviews studies
that have shown shifts in the timing of flowering in plants and
of migration in birds in response to climate warming. The dis-
cussion complements the earlier discussion of shifts in species
ranges in chapter 10 by demonstrating that climate warming is
not just inducing organisms to move in response to global warm-
ing but also adjusting their life histories.
In chapter 13, the Lotka-Volterra equations have been
modified from previous editions to make them more standard,
less cluttered, and easier for students to follow, which is essen-
tial, since these equations are the foundation of the mathematical
ecology covered in the text.
In chapter 14, we revisit predator functional responses
first introduced in chapter 7 by evaluating alternatives to those
models. The Lotka-Volterra models of predator-prey interactions
published in the early twentieth century stimulated a long line
of research. More recently, researchers have offered alternatives
that help identify where those classical mathematical models,
with their simplifying assumptions, apply and where alternative
formulations better account for aspects of predator-prey inter-
actions, particularly at larger spatial and longer temporal
scales. The discussion in this chapter reviews how recent ratio-
dependent functional response models better predict predator

functional responses in experimental and natural settings. The
discussion helps to dispel the idea that mathematical ecology
ceased to develop in the mid-twentieth century and reinforces the
complementary roles of theoretical, experimental, and observa-
tional studies.
In chapter 18, a new concept connects primary producer
diversity to higher levels of primary production. The chapter also
includes a new concept featuring the relationship between levels
of primary production and secondary production. This discussion
provides a basis for introducing the fundamentals of secondary
production. This addition also revisits the ratio-dependent func-
tional responses introduced in chapter 14 by extending the impli-
cations of those models beyond predator functional response to
the trophic structure of ecosystems. The treatment also formally
introduces secondary production, filling a conceptual gap in pre-
vious editions.
In chapter 20, the fields of ecological restoration and
restoration ecology are introduced for the first time. Human
impact on the environment has altered ecological communities
and ecosystems in nearly every corner of the planet. Restoring
Preface xvii
structure and function to these systems emerges as one of the
great contemporary ecological challenges. Increasingly ecolo-
gists addressing this challenge are turning to the conceptual
framework of ecological succession to guide their work. Exam-
ples of such work are included in this chapter to help bridge
the historical divide between ecological theory and restoration
practice.
In chapter 23, the discussion of the Antarctic ozone hole
has been updated to 2013, including 35 years of data from NASA
on the size of the ozone hole. The pattern shows that the maxi-
mum size of the Antarctic ozone hole has stabilized, signaling
a basis for ozone recovery predicted by atmospheric scientists
over the next 50 years, providing a bit of good planetary news.
The growing body of climate change research, published since
the earlier editions of Ecology Concepts and Applications, has
greatly improved understanding of how earth’s changing climate
will impact ecosystems and human populations, if not stabilized.
A discussion of these impacts concludes this edition, underscor-
ing the relevance of ecological knowledge to sustaining natural
as well as human-centered systems.
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xix
xx Preface
Annual Editions: Environment 2015
by Eathorne
ISBN 978-1-25-916115-5
Annual Editions is a compilation of current articles from the
best of the public press. The selections explore the global
environment, the world’s energy, the biosphere, natural
resources, and pollution. Available through Create.
Taking Sides: Clashing Views on
Environmental Issues,
Sixteenth Edition by Easton
ISBN: 978-1-25-916113-1
Taking Sides presents current contro-
versial issues in a debate-style format
designed to stimulate student interest
and develop critical thinking skills.
Each issue is thoughtfully framed with
an issue summary, an issue introduction, and a postscript or
challenge questions. An online Instructor’s Resource Guide
with testing material is available. Available through Create.
Classic Edition Sources: Environmental Studies
Fourth Edition by Thomas Easton
ISBN 978-0-07-352764-2
Sources brings together selections of enduring intellectual
value—classic articles, book excerpts, and research studies—
that have shaped ecology and environmental science. Edited
for length and level, the selections are organized topically.
An annotated table of contents provides a quick and easy
review of the selections. Supported by an online instructor’s
Resource Guide that provides a complete synopsis of each selec-
tion, guidelines for discussing the selection in class, and testing
materials. Available through Create.
Ecology Laboratory Manual, by Vodopich
(ISBN: 978-0-07-338318-7;
MHID: 0-07-338318-X)
Darrell Vodopich, co-author of Biology Laboratory Manual,
has written a new lab manual for ecology. This lab manual
offers straightforward procedures that are doable in a broad
range of classroom, lab, and field situations. The procedures
have specific instructions that can be taught by a teaching
assistant with minimal experience as well as by a professor.
Student Atlas of Environmental
Issues, by Allen
(ISBN: 978-0-69-736520-0;
MHID: 0-69-736520-4)
This atlas is an invaluable pedagogical
tool for exploring the human impact on
the air, waters, biosphere, and land in
every major world region. This infor-
mative resource provides a unique
combination of maps and data that help students understand
the dimensions of the world’s environmental problems and
the geographic basis of these problems.
Acknowledgments
A complete list of the people who have helped me with this
project would be impossibly long. However, during the devel-
opment of this seventh edition, several colleagues freely
shared their ideas and expertise, reviewed new sections, or
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,
Will Pockman, Steve Poe, Bob Sinsabaugh, Alain Thomas,
Tom Turner, Lawrence Walker, Chris Witt, Blair Wolf. I wish
to offer special thanks to Roger Arditi and Lev Ginzburg
for their time and patience in helping me develop sections
on ratio-dependent models of functional response and their
potential contributions to better understanding of predator-
prey interactions and the trophic structure of ecosystems. I am
also grateful to Art Benke for helping me develop an over-
view of secondary production for this edition and for helping
integrate it with discussion of the effects of enrichment on
ecosystem trophic structure. John and Leah Vucetich helped
bring their long-term research on wolf-moose interactions on
Isle Royale to life by graciously allowing use of one of their
many photos of interactions in this model predator and prey
system. In addition, I am indebted to the many students and
instructors who have helped by contacting me with questions
and suggestions for improvements.
I also wish to acknowledge the skillful guidance and work
throughout the publishing process given by many profession-
als associated with McGraw-Hill during this project, including
Becky Olson, Patrick Reidy, Carrie Burger, Fran Simon, April
Southwood, Lynn Breithaupt, Mary Reeg, Angie Sigwarth, Tara
McDermott, and Sheila Frank.
Finally, I wish to thank all my family for support given
throughout the project, especially Paulette Dompeling, Mary Ann
Esparza, Dan Esparza, Hani Molles, Anders Molles, Mary Anne
Nelson, and Keena.
I gratefully acknowledge the many reviewers who, over the
course of the last several revisions, have given of their time and
expertise to help this textbook evolve to its present seventh edi-
tion. Their depth and breadth of knowledge and experience, both
as researchers and teachers, are humbling. They continue my
education, for which I am grateful, and I honestly could not have
continued the improvement of this textbook without them.
I gratefully acknowledge the many reviewers who, over
the course of the last several revisions, have given of their
time and expertise to help this textbook evolve to its present
edition. Their depth and breadth of knowledge and experi-
ence, both as researchers and teachers, are humbling. They
continue my education, for which I am grateful, and I hon-
estly could not have continued the improvement of this text-
book without them.
Reviewers for the Seventh Edition
John Bacheller Hillsborough Community College
Isaac Barjis City University of New York
Dena Berg Tarrant County College NW
Earl R. Beyer Harrisburg Area Community College

Jamal Bittar The University of Toledo
Linda Bruslind Oregon State University
Sherri L. Buerdsell West Virginia Northern Community College
Carrie E. Burdzinski Delta College (University Center, Michigan)
William Dew Nipissing University
Harry G. Deneer University of Saskatchewan
Phil Denette Delgado Community College
Jessica A. DiGirolamo Broward College, Davie, Florida
Angela M. Edwards Trident Technical College
Elyce Ervin University of Toledo
Teresa G. Fischer Indian River State College
Christina Gan Highline Community College
Kathryn Germain Southwest Tennessee Community College
Linda Girouard Brescia University
Judy Gnarpe University of Alberta
Amy D. Goode Illinois Central College
Robert C. Hairston Harrisburg Area Community College
Nasreen S. Haque City University of New York, New York
Daniel P. Herman University of Wisconsin—Eau Claire
Ingrid Herrmann Santa Fe College
Sheela S. Huddle Harrisburg Area Community College
Chike Igboechi Medgar Evers College of the City University
of New York
Ilko G. Iliev Southern University at Shreveport
Debra W. Jackson University of Louisiana at Monroe
John C. Jones Calhoun Community College
Judy Kaufman Monroe Community College
Peter S. Kourtev Central Michigan University
Jonathan N. Lawson Collin College, Plano Texas
Suzanne Long Monroe Community College
Mary Ann Merz West Virginia Northern Community College
Matthew Morgan Greenville Technical College
Christian Nwamba Wayne County Community
College District
Amanda Thigpen Parker Pearl River Community College
Marceau Ratard Delgado Community College
Geraldine H. Rimstidt Daytona State College
Seth Ririe Brigham Young University—ldaho
David M. Rollins University of Maryland, College Park &
Prince Georges Community College
Ben Rowley University of Central Arkansas
Eleftherios “Terry” Saropoulos Vanier College
Arif Sheena MacEwan College, Alberta, Canada
Richard H. Shippee Vincennes University
Sasha A. Showsh University of Wisconsin—Eau Claire
Susan J. Stamler College of DuPage
Ronald J. Stewart Humber ITAL, Toronto, Ontario
Victoria Auerbuch Stone UC Santa Cruz
David J. Wartell Harrisburg Area Community College
TitYee Wong University of Memphis
Reviewers for the Sixth Edition
Michael Henshaw Grand Valley State University
Thomas Nash Arizona State University
Thomas Schoener University of California—Davis
Kevin Woo University of Central Florida
Deborah Waller Old Dominion University
William Kroll Loyola University of Chicago
James Manhart Texas A&M University
Jonathan Benstead University of Alabama
Robert Sanders Temple University
Preface xxi
Jerry Baskin University of Kentucky
Thomas O. Crist Miami University
Peter Alpert University of Massachusetts—Amherst
Mark Pyron Ball State University
Mary Bremigan Michigan State University
Reviewers for the Fifth Edition
Joel S. Brown University of Illinois—Chicago
Peter E. Busher Boston University
Lloyd Fitzpatrick University of North Texas
James A. Fordyce University of Tennessee
David L. Gorchov Miami University
Jamie Kneitel California State University—Sacramento
John C. Krenetsky Metropolitan State College of Denver
Amy E. Lesen Pratt Institute
D. Nicholas McLetchie University of Kentucky
Thomas Pliske Florida International University
Nathan J. Sanders University of Tennessee
Robert M. Schoch Boston University
John F. Weishampel University of Central Florida
Reviewers for the Fourth Edition
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Frank S. Gilliam Marshall University
Colleen Hatfield Rutgers University
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xxii Preface
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Preface xxiii
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Manuel C. Molles Jr.

A yellow-rumped warbler, Dendroica coronata, feeding young.
Ecological studies of warblers have made fundamental contributions
to the growth of ecological understanding.
CHAPTER CONCEPTS
1.1 Ecologists study environmental
relationships ranging from those
of individual organisms to factors
influencing global-scale processes.
Concept 1.1 Review 3
1.2 Ecologists design their studies based on
their research questions, the temporal
and spatial scale of their studies, and
available research tools. 3
Concept 1.2 Review 8
Investigating the Evidence 1:
The Scientific Method—Questions
and Hypotheses 9
Summary 10
Key Terms 10
Review Questions 10
Introduction
to Ecology
1
Historical Foundations
and Developing Frontiers
LEARNING OUTCOME
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-
ships between organisms and the environment,
has been a focus for human study for as long as
we have existed as a species. Our survival has depended upon
how well we could observe variations in the environment and
predict the responses of organisms to those variations. The
2 earliest hunters and gatherers had to know the habits of their
animal prey and where to find food plants. Later, agricultur-
ists had to be aware of variations in weather and soils and of
how such variation might affect crops and livestock.
Today, most of earth’s human population live in cities and
most of us have little direct contact with nature. More than ever
before, though, the future of our species depends on how well we
understand the relationships between organisms and the environ-
ment. Our species is rapidly changing earth’s environment, yet
we do not fully understand the consequences of these changes.
For instance, human activity has increased the quantity of nitro-
gen cycling through the biosphere, changed land cover across
the globe, and increased the atmospheric concentration of CO2.
Changes such as these threaten the diversity of life on earth and
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
2 Chapter 1 Introduction to Ecology

Behind the simple definition of ecology lies a broad sci- Biosphere

entific discipline. Ecologists may study individual organisms,
What role does concentration of
entire forests or lakes, or even the whole earth. The mea- atmospheric CO2 play in the
surements made by ecologists include counts of individual regulation of global
organisms, rates of reproduction, or rates of processes such temperature?
as photosynthesis and decomposition. Ecologists often spend
as much time studying nonbiological components of the
environment, such as temperature or soil chemistry, as they Region
spend studying organisms. Meanwhile, the “environment” of
How has geologic history
organisms in some ecological studies are other species. While influenced regional diversity
you may think of ecologists as typically studying in the field, within certain groups of
some of the most important conceptual advances in ecology organisms?
have come from ecologists who build theoretical models or
do ecological research in the laboratory. Clearly, our simple
Landscape
definition of ecology does not communicate the great breadth
of the discipline or the diversity of its practitioners. To get a How do vegetated corridors
affect the rate of movement by
better idea of what ecology is, let’s briefly review the scope mammals among isolated forest
of the discipline. fragments?

1.1 Overview of Ecology

Ecosystem
LEARNING OUTCOMES
How does fire affect nutrient
After studying this section you should be able to do the following: availability in grassland
ecosystems?
1.2 Describe the levels of ecological organization, for
example, population, studied by ecologists.
1.3 Distinguish between the types of questions
addressed by ecologists working at different levels of Community
organization.
What factors influence the
1.4 Explain how knowledge of one level of ecological number of large mammal
organization can help guide research at another species living together in
level of organization. African grasslands?

Ecologists study environmental relationships ranging Interactions

from those of individual organisms to factors influencing
global-scale processes. This broad range of subjects can Do predators influence where
zebras feed in the landscape?
be organized by arranging them as levels in a hierarchy of
ecological organization, such as that imbedded in the brief
table of contents and the sections of this book. Figure 1.1
attempts to display such a hierarchy graphically.
Historically, the ecology of individuals, which is presented Population
at the base of figure 1.1, has been the domain of physiological What factors control zebra
ecology and behavioral ecology. Physiological ecologists have populations?
emphasized the evolution (a process by which populations
change over time) of physiological and anatomical mechanisms
by which organisms solve problems posed by physical and
chemical variation in the environment. Meanwhile, behavioral
ecologists have focused principally on evolution of behaviors Individuals
that allow animals to survive and reproduce in the face of envi- How do zebras regulate their
ronmental variation. Physiological and behavioral ecology are internal water balance?
informed by evolutionary theory, as are all other areas of ecology.
There is a strong conceptual linkage between ecological
studies of individuals and of populations particularly where
they concern evolutionary processes. Population ecology is Figure 1.1 Levels of ecological organization and examples of the
centered on the factors influencing population structure and kinds of questions asked by ecologists working at each level. These
process, where a population is a group of individuals of a sin- ecological levels correspond broadly to the sections of this book.
gle species inhabiting a defined area. The processes studied
by population ecologists include adaptation, extinction, the
 Chapter 1 Introduction to Ecology 3

distribution and abundance of species, population growth and

regulation, and variation in the reproductive ecology of spe-
cies. Population ecologists are particularly interested in how
these processes are influenced by nonbiological and biologi-
cal components of the environment.
Bringing biological components of the environment into
the picture takes us to the next level of organization, the ecol-
ogy of interactions such as predation, parasitism, and com-
petition. Ecologists who study interactions between species
have often emphasized the evolutionary effects of the inter-
action on the species involved. Other approaches explore the
effect of interactions on population structure or on properties
of ecological communities.
The definition of an ecological community as an associa-
tion of interacting species links community ecology with the
ecology of interactions. Community and ecosystem ecology (a)
have a great deal in common, since both are concerned with the
factors controlling multispecies systems. However, the objects
of their study differ. While community ecologists concentrate on
the organisms inhabiting an area, ecosystem ecologists include
the physical and chemical factors influencing the community
and focus on processes such as energy flow and decomposition.
To simplify their studies, ecologists have long attempted
to identify and study isolated communities and ecosystems.
However, all communities and ecosystems on earth are open
systems subject to exchanges of materials, energy, and organ-
isms with other communities and ecosystems. The study of
these exchanges, especially among ecosystems, is the intel-
lectual territory of landscape ecology. However, landscapes
are not isolated either but part of geographical regions sub-
ject to large-scale and long-term regional processes. These
regional processes are the subjects of geographic ecology.
Geographic ecology in turn leads us to the largest spatial scale (b)
and highest level of ecological organization—the biosphere, Figure 1.2 Two rapidly developing frontiers in ecology.
the portions of the earth that support life, including the land, (a) Aeroecology: the interdisciplinary study of the ecology of the earth-
waters, and atmosphere. atmosphere boundary (Kunz et al. 2008). New tools, such as the Indigo/
While this description of ecology provides a brief preview FLIR Merlin mid thermal camera that took this thermal infrared image
of flying Brazilian free-tailed bats, Tadarida braziliensis, have opened
of the material covered in this book, it is a rough sketch and this developing frontier in ecology. This image depicts variation in the
highly abstract. To move beyond the abstraction represented surface temperature of these bats. Thermal infrared technology makes
by figure 1.1, we need to connect it to the work of the scien- it possible not only to detect and record the presence of free-ranging
tists who have created the discipline of ecology. To do so, let’s nocturnal organisms, but also to investigate their physiology and ecology
briefly review the research of ecologists working at a broad in a noninvasive manner (see chapter 5, p. 114). (b) Urban ecology: the
study of urban areas as complex, dynamic ecological systems, influenced
range of ecological levels emphasizing links between histori- by interconnected, biological, physical, and social components. As ecolo-
cal foundations and some developing frontiers (fig. 1.2). gists focus their research on the environment where most members of our
species live, they have made unexpected discoveries about the ecology of
urban centers such as the city of Baltimore (see chapter 19, p. 432).

Concept 1.1 Review

1. How does the level of ecological organization an ecolo-
gist studies influence the questions he or she poses?
2. While an ecologist may focus on a particular level of
ecological organization shown in figure 1.1, might other
levels of organization be relevant, for example, does an
ecologist studying factors limiting numbers in a popula-
tion of zebras need to consider the influences of interac-
tions with other species or the influences of food on the
survival of individuals?
1.2 Sampling Ecological Research
LEARNING OUTCOMES
After studying this section you should be able to do the following:
1.5 Describe some emerging frontiers in ecology.
1.6 Explain how the use of stable isotopes has extended
what it is possible to know about the ecology of warblers.
1.7 Compare the spatial and temporal scales addressed
by the research of Robert MacArthur, Nalini
Nadkarni, and Margaret Davis.
4 Chapter 1 Introduction to Ecology
Ecologists design their studies based on their research
questions, the temporal and spatial scale of their studies, and
available research tools. Because the discipline is so broad,
ecological research can draw from all the physical and biologi-
cal sciences. The following section of this chapter provides a
sample of ecological questions and approaches to research.
The Ecology of Forest Birds:
Old Tools and New
Robert MacArthur gazed intently through his binoculars. He
was watching a small bird, called a warbler, searching for
insects in the top of a spruce tree. To the casual observer it might
have seemed that MacArthur was a weekend bird-watcher. Yes,
he was intensely interested in the birds he was watching, but he
was just as interested in testing ecological theory.
The year was 1955, and MacArthur was studying the
ecology of five species of warblers that live together in the
spruce forests of northeastern North America. All five warbler
species, Cape May (Dendroica tigrina), yellow-rumped
(D. coronata), black-throated green (D. virens), blackbur-
nian (D. fusca), and bay-breasted (D. castanea), are about
the same size and shape and all feed on insects. Theory pre-
dicted that two species with identical ecological requirements
would compete with each other and that, as a consequence,
they could not live in the same environment indefinitely. Mac-
Arthur wanted to understand how several warbler species with
apparently similar ecological requirements could live together
in the same forest.
The warblers fed mainly by gleaning insects from the
bark and foliage of trees. MacArthur predicted that these
Cape May warbler
New needles and New needles and buds
buds at top of tree of upper branches
Bay-breasted
warbler
Old needles and bare
and lichen-covered
middle branches
Figure 1.3 Warbler feeding zones shown in beige. The several warbler species that coexist in the forests of northeastern North America feed in
distinctive zones within forest trees.
warblers might be able to coexist and not compete with each
other if they fed on the insects living in different zones within
trees. To map where the warblers fed, he subdivided trees into
vertical and horizontal zones. He then carefully recorded the
amount of time warblers spent feeding in each.
MacArthur’s prediction proved to be correct. His quan-
titative observations demonstrated that the five warbler spe-
cies in his study area fed in different zones in spruce trees. As
figure 1.3 shows, the Cape May warbler fed mainly among
new needles and buds at the tops of trees. The feeding zone
of the blackburnian warbler overlapped broadly with that of
the Cape May warbler but extended farther down the tree. The
black-throated green warbler fed toward the trees’ interiors.
The bay-breasted warbler concentrated its feeding in the inte-
rior of trees. Finally, the yellow-rumped warbler fed mostly
on the ground and low in the trees. MacArthur’s observations
showed that though these warblers live in the same forest,
they extract food from different parts of that forest. He con-
cluded that feeding in different zones may reduce competition
among the warblers of spruce forests.
MacArthur’s study (1958) of foraging by warblers is
a true classic in the history of ecology. However, like most
studies it raised as many questions as it answered. Scientific
research is important both for what it teaches us directly about
nature and for how it stimulates other studies that improve
our understanding. MacArthur’s work stimulated numerous
studies of competition among many groups of organisms,
including warblers. Some of these studies produced results
that supported his work and others produced different results.
All added to our knowledge of competition between species
and of warbler ecology.
Blackburnian warbler Black-throated
green warbler
New needles and buds
and some older needles
Yellow-rumped
warbler
Bare or lichen-covered lower
trunk and middle branches

Figure 1.4 A male American redstart, Setophaga ruticilla.
Mature male American redstarts are highly territorial, dominating
high-quality feeding territories in their tropical wintering grounds,
pushing most female redstarts and young males into poorer-quality
feeding habitats.
Nearly half a century after Robert MacArthur studied the
feeding ecology of warblers through the lenses of his binocu-
lars, a team of Canadian and U.S. scientists led by Ryan Norris
(Norris et al. 2005) worked to develop tools capable of pen-
etrating the feeding habitats of wide-ranging migratory birds.
The object of their study was the American redstart (Setophaga
ruticilla), another colorful member of the warbler family Paru-
lidae (fig. 1.4). American redstarts, like the warblers studied by
MacArthur, are long-distance migrants, nesting in temperate
North America but spending their winters mainly in tropical Cen-
tral America, northern South America, and the Caribbean islands.
Historically, studies of wide-ranging bird species, such
as the American redstart, have focused mainly on their tem-
perate breeding grounds. However, observations by ecolo-
gists had long suggested that the success of an individual
migratory bird during the breeding season may depend criti-
cally on the environmental conditions it experienced on its
tropical wintering grounds. For example, it has been well
established that male migratory birds, arriving early on the
breeding grounds, are generally in better physical condition
compared to those arriving later. Early arrivals also gen-
erally obtain the best breeding territories and have higher
reproductive success.
Variation in arrival times and physical condition led ecol-
ogists to ponder the connection between events on the win-
tering grounds and subsequent reproductive success among
birds in their breeding habitats. To answer such a question,
we need a great deal of information, including where indi-
vidual birds live on the wintering grounds, how the winter
habitat correlates with physical condition during migration,
how winter habitat influences time of arrival on the breeding
grounds, and whether winter habitat correlates with reproduc-
tive success on the breeding grounds. Clearly, the amount of
information required to answer such questions, concerning
environments separated by thousands of kilometers (fig. 1.5),
exceeds what one person, or even a large team, can learn
through the lenses of binoculars.
Chapter 1 Introduction to Ecology 5
American redstarts breed across
much of North America, preferring
forests with abundant shrubs.
American redstarts winter
mainly on the Caribbean
islands and the surrounding
mainland.
Breeding grounds
Wintering grounds
Figure 1.5 Map of the breeding and wintering grounds of the
American redstart, Setophaga ruticilla.
Often, ecologists have pioneered the use of more power-
ful research tools, as the complexity of their questions have
increased. A tool to which ecologists turn increasingly to
understand the ecology of migratory birds is stable isotope
analysis (see chapter 6, p. 145). Isotopes of a chemical ele-
ment, for example, isotopes of carbon, have different atomic
masses as a result of having different numbers of neutrons.
Carbon, for instance, has three isotopes (listed in order of
increasing mass): 12C, 13C, and 14C. Of these three, 12C and
13
C are stable isotopes because they do not undergo radioac-
tive decay, whereas 14C decays radioactively and is therefore
unstable. Stable isotopes have proven useful in the study of
ecological processes—for example, identifying food sources,
because the proportions of various isotopes differ across the
environment.
Stable isotope analysis provides ecologists with a new
type of “lens” capable of revealing ecological relationships
that would otherwise remain invisible. For example, ecolo-
gists using stable isotope analysis can track habitat use by
American redstarts on their wintering grounds. In Jamaica,
older male American redstarts, along with some females,
spend the winter in higher-productivity mangrove forest hab-
itats, pushing most females and younger males into poorer-
quality, dry scrub habitat. The dominant plants in these two
habitats and the insects that feed on them contain different
proportions of the carbon isotopes 12C and 13C. Therefore,
the tissues of the birds spending their winters in the produc-
tive mangrove habitat (lower 13C) and those spending the
winters in the poor scrub habitat (higher 13C) are in effect
chemically tagged. As a consequence, today’s ecologist can
analyze a very small sample of blood from an American red-
start when it arrives on its temperate breeding ground and
6 Chapter 1 Introduction to Ecology

know the habitat where it spent the winter. When Ryan Nor-
ris and his research team made such measurements, they
found that male redstarts that had spent the winter in the
more productive mangrove habitat arrived on the breeding
grounds earlier and produced significantly more young birds
that survived to fledging.
Stable isotope analysis and the role that it has played in
elucidating the ecology of a diversity of organisms will thread
its way through the text. As is often the case in science, new
tools create new research frontiers. Another of those frontiers
is to be found in the canopies of forests.

Forest Canopy Research:

A Physical and Scientific Frontier
Studies of warblers showcase how ecologists approach
studies of one or a few species. Other ecologists have been
concerned with the ecology of entire forests, lakes, or
grasslands, which they treat as ecosystems. An ecosystem
includes all the organisms that live in an area and the physi-
cal environment with which those organisms interact. Many
ecosystem studies have focused on nutrients, the raw mate-
rials that an organism must acquire from the environment
to live.
For ecologists who study the budgets of nutrients such as
nitrogen, phosphorus, or calcium, one of the first steps is to
inventory their distribution within an ecosystem. Inventories
by Nalini Nadkarni (1981, 1984a, 1984b) changed our ideas
of how tropical and temperate rain forests are structured
and how they function. With the aid of mountain-climbing
equipment, Nadkarni slowly made her first ascent into the
canopy of the Costa Rican rain forest, a world explored by
few others and where she was to become a pioneer (fig. 1.6).
She stood on the rain forest floor and wondered about the
diversity of organisms and ecological relationships that
might be hidden in the canopy high above. Her wonder soon
gave way to determination, and Nadkarni not only visited
the canopy but was among the first to explore the ecology of
this unseen world.
Because of leaching by heavy rains, many rain forest
soils are poor in nutrients such as nitrogen and phosphorus.
The low availability of nutrients in many rain forest soils has
produced one of ecology’s puzzles. How can the prodigious
life of rain forests be maintained on such nutrient-poor soils?
Many factors contribute to the maintenance of this intense
biological activity. Nadkarni’s research in the treetops uncov-
ered one of those factors, a significant store of nutrients in the
rain forest canopy.
The nutrient stores in the rain forest canopy are associ-
ated with epiphytes. Epiphytes are plants, such as many
orchids and ferns, that live on the branches and trunks of other
plants. Epiphytes are not parasitic: they do not derive their
nutrition from the plant they grow on. As they grow on the
branches of a tree they begin to trap organic matter, which
eventually forms a mat. Epiphyte mats increase in thickness
up to 30 cm, providing a complex structure that supports a
diverse community of plants and animals.
Figure 1.6 Exploring the rain forest canopy. What Nalini Nadkarni
discovered helped solve an ecological puzzle.
Epiphyte mats contain significant quantities of nutrients.
Nadkarni estimated that these quantities in some tropical rain
forests are equal to about half the nutrient content of the foli-
age of the canopy trees. In the temperate rain forests of the
Olympic Peninsula in Washington, the mass of epiphytes is
four times the mass of leaves on their host trees.
Nadkarni’s research showed that in both temperate and
tropical rain forests, trees access these nutrient stores by send-
ing out roots from their trunks and branches high above the
ground. These roots grow into the epiphyte mats and extract
nutrients from them. As a consequence of this research, we
now know that to understand the nutrient economy of rain for-
ests the ecologist must venture into the treetops.
Easier means of working in the rain forest canopy have
been developed, and this research is no longer limited to the
adventurous and agile. New ways to access the forest canopy
range from hot air balloons and aerial trams to large cranes.
The Wind River Canopy Crane offers scientists access to any
level within a 70 m tall coniferous forest in a 2.3 ha area near
the Columbia River Gorge in Washington (fig. 1.7). Research
projects supported—and made far easier—by this crane and
others have included the ecology of migratory birds in the
forest canopy, photosynthesis by epiphytes living at differ-
ent canopy heights, and vertical stratification of habitat use
by bats and beetles (Ozanne et al. 2003). By 2006, there were
12 canopy cranes facilitating canopy research in temperate
and tropical forests worldwide (Stork 2007). Nadkarni points

Gondola housing scientists can
Canopy zonation be lowered to study any level in
the canopy.
Over 40 m:
Physical conditions: greatest
exposure to sunlight and winds, Height (m)
highest variability in temperature
Characteristic animals: red crossbill,
warblers, flying squirrel
75
15 to 40 m 60
Physical conditions: partial shading,
lower exposure to winds, more equable
temperatures 45
Characteristic animals: chickadees,
nuthatches, varied thrush
30
Ground to 15 m 15
Physical conditions: lowest light
intensity and reduced temperature
variation, diminished wind 0
Characteristic animals: towhees, Douglas
American robin, winter wren, black- fir
tailed deer, coyote
Figure 1.7 The Wind River Canopy Crane provides access to the forest canopy for a broad range of ecology and ecological studies.
out, in response to these developments, that the canopy as a
physical frontier may be closing, but its exploration as a sci-
entific frontier is just beginning, particularly as we attempt to
predict the ecological consequences of climate change.
Climatic and Ecological Change:
Past and Future
The earth and its life are always changing. However, many
of the most important changes occur over such long periods
of time or at such large spatial scales that they are difficult
to study. Two approaches that provide insights into long-term
and large-scale processes are studies of pollen preserved in
lake sediments and evolutionary studies.
Margaret Davis (1983, 1989) carefully searched through
a sample of lake sediments for pollen. The sediments had
come from a lake in the Appalachian Mountains, and the
pollen they contained would help her document changes in
the community of plants living near the lake during the past
several thousand years. Davis is a paleoecologist trained to
think at very large spatial scales and over very long periods of
time. She has spent much of her professional career studying
changes in the distributions of plants during the Quaternary
period, particularly during the most recent 20,000 years.
Some of the pollen produced by plants that live near a
lake falls on the lake surface, sinks, and becomes trapped in
Chapter 1 Introduction to Ecology 7
Crane operator can swing crane
arm 360˚ and position gondola
at any distance along crane arm.
Pacific Grand Pacific Western Pacific Western
silver fir fir yew hemlock dogwood red cedar
Tree species
lake sediments. As lake sediments build up over the centu-
ries, this pollen is preserved and forms a historical record
of the kinds of plants that lived nearby. As the lakeside veg-
etation changes, the mix of pollen preserved in the lake’s
sediments also changes. In the example shown in figure 1.8,
pollen from spruce trees, Picea spp., first appears in lake
sediments about 12,000 years ago then pollen from beech,
Fagus grandifolia, occurs in the sediments beginning
about 8,000 years ago. Chestnut pollen does not appear
in the sediments until about 2,000 years ago. The pollen
from all three tree species continues in the sediment record
until about 1920, when chestnut blight killed most of the
chestnut trees in the vicinity of the lake. Thus, the pollen
preserved in the sediments of lakes can be used to recon-
struct the history of vegetation in the area. Margaret B.
Davis, Ruth G. Shaw, and Julie R. Etterson review extensive
evidence that during climate change, plants evolve, as well
as disperse (Davis and Shaw 2001; Davis, Shaw, and Etter-
son 2005). As climate changes, plant populations simultane-
ously change their geographic distributions and undergo the
evolutionary process of adaptation, which increases their
ability to live in the new climatic regime. Meanwhile, evi-
dence of evolutionary responses to climate change is being
discovered among many animal groups. William Bradshaw
and Christina Holzapfel (2006) summarized several stud-
ies documenting evolutionary change in northern animals,
8 Chapter 1 Introduction to Ecology
Lake profile
Sediments
Figure 1.8 The vegetation history of landscapes can be reconstructed using the pollen contained within the sediments of nearby lakes.
ranging from small mammals and birds to insects (fig. 1.9),
in response to increasing growing season length as a conse-
quence of the now-well-documented phenomenon of global
Figure 1.9 Studies indicate that north American red squirrels,
Tamiasciurus hudsonicus, have been undergoing rapid evolution for
earlier breeding, during a recent period of increased average spring
temperatures in Canada’s Yukon Territory (Réale et al. 2003).
Chestnut decline Present Pollen
due to blight 100
Chestnut
2,000
Years
Beech
8,000
12,000 Spruce
Sediment core
warming (see chapter 23, p. 519). Research such as that
by Davis and her colleagues will be essential to predicting
and understanding ecological responses to global climate
change.
In the remainder of this book we will fill in the details
of the sketch of ecology presented in this chapter. This brief
survey has only hinted at the conceptual basis for the research
described. Throughout this book we emphasize the concep-
tual foundations of ecology. Each chapter focuses on a few
ecological concepts. We also explore some of the applications
associated with the concepts introduced. Of course, the most
important conceptual tool used by ecologists is the scientific
method, which is introduced on page 9.
We continue our exploration of ecology in section I with
natural history and evolution. Natural history is the founda-
tion on which ecologists build modern ecology for which evo-
lution provides a conceptual framework. A major premise of
this book is that knowledge of natural history and evolution
improves our understanding of ecological relationships.
Concept 1.2 Review
1. How were the warbler studies of Robert MacArthur and
those that focused on the American redstart similar?
How did they differ?
2. What aspects of Nalini Nadkarni’s research identify it
as “ecosystem ecology”? Give examples of research
in forest canopies that would address other levels of
ecological organization (for examples, see fig. 1.1).
3. The discussion of the research by Margaret Davis and
her colleagues did not identify the questions that they
addressed. What research questions can we infer from
the above description of their work?
 Chapter 1 Introduction to Ecology 9

Investigating the Evidence 1

The Scientific Method—Questions and Hypotheses

LEARNING OUTCOMES Information: Testing a hypothesis,

After studying this section you should be able to do the following: Observation whatever the outcome,
Experiment increases the pool of
1.8 Distinguish between questions and hypotheses in the Modeling information.
scientific process. Published Studies
Hypothesis
1.9 Discuss the scientific method, emphasizing hypoth- not supported:
esis testing. Change hypothesis Question
in light of new
information.
Ecologists explore the relationships between organisms and Hypothesis
environment using the methods of science. The series of boxes
called “Investigating the Evidence” that are found throughout Prediction
the chapters of this book discuss various aspects of the sci- Hypothesis
supported:
entific method and its application to ecology. While each box Conduct additional
describes only a small part of science, taken together, they rep- Test of hypothesis:
tests of the
Observation
resent a substantial introduction to the philosophy, techniques, hypothesis.
Experiment
and practice of ecological science. Modeling
Let us begin this distributed discussion with the most
basic point. What is science? The word science comes from a Data:
Latin word meaning “to know.” Broadly speaking, science is Gathering
a way of obtaining knowledge about the natural world using Management
certain formal procedures. Those procedures, which make up Display
Summary Statistics
what we call “the scientific method,” are outlined in figure 1. Statistical Analysis
Despite a great diversity of approaches to doing science, Accept/Reject Hypothesis
sound scientific studies have many methodological charac-
teristics in common. The most universal and critical aspects Figure 1 Graphic summary of the scientific method. The scien-
of the scientific method are: asking interesting questions and tific method centers on the use of information to propose and test
forming testable hypotheses. hypotheses through observation, experiment, and modeling.

Question Hypotheses
Questions and Hypotheses
What do scientists do? Simply put, scientists ask and attempt
to find answers to questions about the natural world. Ques-
tions are the guiding lights of the scientific process. With-
out them, exploration of nature lacks focus and yields little
understanding of the world. Let’s consider a question asked
by an ecologist discussed in this chapter. The main ques-
tion asked by Robert MacArthur in his studies of warblers
(p. 4) was something like the following: “How can several
species of insect-eating warblers live in the same forest with-
out one species eventually excluding the others through com-
petition?” While this focus on questions may seem obvious,
one of the most common questions asked of scientists at semi-
nars and professional meetings is, “What is your question?”
If scientists are in the business of asking questions
about nature, where does a hypothesis enter the process?
A hypothesis is a possible answer to a question. MacArthur’s
main hypothesis (possible answer to his question) was: “Sev-
eral warbler species are able to coexist because each species
feeds on insects living in different zones within trees.”
Once a scientist or team of scientists proposes a hypoth-
esis (or multiple alternative hypotheses), the next step in the
scientific method is to determine its validity by testing predic-
tions that follow from the hypothesis. Three fundamental ways
to test hypotheses are through observation, experiments, and
modeling. These approaches, which are all represented in fig-
ure 1, will be discussed in detail in the “Investigating the Evi-
dence” boxes and in the research discussed in later chapters.
CRITIQUING THE EVIDENCE 1
1. How does the development of new research tools, such as
canopy cranes and stable isotope analysis, affect the pro-
cess of science as outlined by figure 1 of this “Investigat-
ing the Evidence” box?
10 Chapter 1 Introduction to Ecology
Summary
Ecologists study environmental relationships ranging from
those of individual organisms to factors influencing global-
scale processes. The research focus and questions posed by
ecologists differ across the levels of ecological organization
studied.
Ecologists design their studies based on their research
questions, the temporal and spatial scale of their stud-
ies, and available research tools. With this brief review
of research approaches and topics, we return to the question
asked at the beginning of the chapter: What is ecology? Ecol-
ogy is indeed the study of relationships between organisms
and the environment. However, as you can see from the studies
Key Terms
adaptation 7 ecology 1
aeroecology 3 ecosystem 6
biosphere 3 epiphyte 6
Review Questions
1. Faced with the complexity of nature, ecologists have divided the
field of ecology into subdisciplines, each of which focuses on
one of the levels of organization pictured in figure 1.1. What is
the advantage of developing such subdisciplines within ecology?
2. What are the pitfalls of subdividing nature in the way it is repre-
sented in figure 1.1? In what ways does figure 1.1 misrepresent
nature?
3. What could you do to verify that the distinct feeding zones
used by the warblers studied by MacArthur (see fig. 1.3) are
the result of ongoing competition between the different species
of warblers? How might you examine the role of competition
in keeping some American redstarts out of the most productive
feeding areas on their wintering grounds?
4. Although Nalini Nadkarni’s studies of the rain forest canopy
addressed a question related to ecosystem structure, the patterns
reviewed, ecologists study those relationships over a large
range of temporal and spatial scales using a wide variety of
approaches. Ecology includes Davis’s studies of vegetation
moving across the North American continent over a span of
thousands of years. Ecology also includes the observational
studies of birds in contemporary forests by MacArthur. Ecolo-
gists may study processes on plots measured in square centime-
ters or, like those studying the ecology of migratory birds, study
areas may span thousands of kilometers. Important ecological
discoveries have come from Nadkarni’s probing of the rain for-
est canopy and from traces of stable isotopes in a droplet of
blood. Ecology includes all these approaches and many more.
evolution 2 stable isotope analysis 5
nutrient 6 urban ecology 3
of nutrient storage in rain forest canopy resulted from the biol-
ogy of individual organisms, populations of organisms, and
communities of species. Explain.
5. What do the studies of Margaret Davis tell us about the compo-
sition of forests in the Appalachian Mountains during the past
12,000 years (see fig. 1.8)? Based on this research, what pre-
dictions might you make about the future composition of these
forests?
6. During the course of the studies reviewed in this chapter, each
scientist or team of scientists measured certain variables. What
major variable studied by Margaret Davis and her research
team distinguishes their work from that of the other research
reviewed in the chapter?
Another random document with
no related content on Scribd:
 Fig. 42. A transsection through the posterior region of the œsophagus of the
hibernating animal, under low magnification; e, epithelium; cm, circular muscles;
lm, longitudinal muscles; mm, muscularis mucosa; sm, submucosa; s, serosa.

In the posterior region of the œsophagus, as may be seen by

comparison of figures 41 and 42, the wall as a whole is about one
third thicker than in the anterior region just described, though how
much of this difference is due to different degrees of distension or
contraction it is hard to say.
The epithelium, e, is in the tissue studied thrown into less
complicated folds than in the anterior region, and is not so thick.
The submucosa, sm, if the entire layer may be so called, has
about the same thickness and structure as in the more anterior
region; but instead of the small and widely scattered bundles of
longitudinal muscle fibers there is a distinct layer of muscle which
may be called the muscularis mucosa, mm, lying about midway
between the epithelium and the circular muscle layer.
The muscularis mucosa is somewhat variable in thickness and is
thrown into folds that correspond to the larger folds of the epithelium
and the submucosa; one of these folds is shown in Figure 42. The
fibers of the muscularis mucosa are apparently all longitudinal in
position.
Outside of the submucosa is a layer of circular muscle fibers, cm;
it is here somewhat wider and more dense than in the anterior
region.
The longitudinal muscle layer (Fig. 42, lm) is much wider and more
compact than in the anterior region. The fibers are indistinctly divided
into large irregular masses as shown in the figure.
The serosa (Fig. 42, s) is a varying but fairly thick layer that is
quite distinct from the longitudinal muscle layer. It consists of the
usual connective tissue groundwork with scattered blood-vessels.
The epithelium, as was said above, is thicker and somewhat more
folded in the anterior than in the posterior region, and in the former
region is partially ciliated while in the latter cilia are entirely wanting.
With these exceptions the epithelium is practically the same in the
two regions.
 Figure 43 represents the epithelium from the anterior region as
seen under high magnification. The outlines of all the cells could not
be determined but if each nucleus represents a cell there are twenty-
five or thirty layers of cells. The nuclei are arranged in two dense,
irregular groups, one along the base of the epithelium, the other
about two thirds of the distance from the base to the free border. The
basal nuclei are perhaps slightly larger and more rounded than those
of the distal group. Between these two groups are numerous more
scattered nuclei; while scattered through the epithelium, except near
the free border, are smaller, round nuclei that stain somewhat darker
than the rest; these, from their size and appearance, seem possibly
to belong to an invisible network of connective tissue that has
penetrated the epithelium from the surrounding mucosa.
 Fig. 43. The epithelium of the anterior region of the œsophagus of the
hibernating animal, under high magnification.

The free border of the epithelium consists of long, ciliated,

columnar cells in which the cell walls may be easily seen. The cilia
are of average length and even in this anterior region are not
everywhere present; possibly they are arranged in bands, but the
material at hand was not sufficient to determine this. As was noted
above, cilia are wanting in the posterior region.
The only differences noted in the anterior region of the
œsophagus between the feeding and the hibernating conditions are
in the muscularis mucosa and the epithelium. As was noted above,
the muscularis mucosa is practically absent in the hibernating stage,
being represented only by a few small, scattered bundles of
longitudinal muscle fibers; while in the feeding stage there is a
narrow but fairly distinct layer to represent the muscularis mucosa.
 Fig. 44. The epithelium of the anterior region of the œsophagus of
the feeding animal, under high magnification.

The difference in the appearance of the epithelium is not striking.

The nuclei are somewhat larger in the feeding stage and, instead of
being crowded into a basal and a median zone, as noted in the
hibernating conditions, they form a dense basal zone, but show no
indication of medial zone. From the dense basal zone the nuclei
become more scattered towards the free surface and are rarely
found closer to the surface than is shown in Figure 44. The smaller
nuclei scattered among the larger ones, noted in connection with the
hibernating stage, are not here seen.
As in the hibernating stage cilia are present on some but not all
cells of this region.
The only noticeable difference between the feeding and
hibernating conditions of the posterior region of the œsophagus is in
the epithelium, which, as in the feeding condition of the anterior
œsophagus, exhibits but one zone of closely set nuclei, that at the
base of the epithelium.
The Stomach. The stomach was sectioned in three regions, as
shown in Figure 35: (1) in the cardiac region very near the opening
of the œsophagus; (2) in the middle or fundic region; and (3) in the
region near the opening of the pylorus. The first two sections are in
the first or large region of the stomach; the third section is in the
second or small region of the stomach (Fig. 35).
The wall as a whole is thickest in the fundus, being there
practically twice as thick as in the pyloric and half again as thick as in
the cardiac region. This great thickening is due mainly to a
thickening of the middle or oblique layer of muscle, which is here
remarkably developed. The mucosa is of nearly uniform thickness in
the different regions and will be described later.
Since there is no striking difference beside that of thickness in the
general structure of the wall of the different regions, the pyloric
region, as seen under low magnification, will now be described (Fig.
45).
 The mucosa, m, consists of fairly long glands underlaid by a well-
marked muscularis mucosa, mm, the latter exhibiting a compact
circular layer over a wider but more scattered layer of longitudinal
fibers. A considerable amount of fibrous connective tissue lies
among the muscle fibers. The circular layer of the muscularis
mucosa sends towards the surface numerous strands or septa
between the glands; six or eight of these are seen in the figure.
These strands are not nearly so numerous in the large region of the
stomach. As was said, the outer or longitudinal layer of the
muscularis mucosa is wider but less compact than the circular and
its bundles of fibers are seen in the figure as a layer of large,
scattered dots just beneath the circular layer.
The submucosa, sm, is of average thickness and density. In the
fundic and cardiac regions it seems to extend between the circular
and oblique layers; at any rate, there is a considerable layer of
connective tissue between these two muscular layers.
The circular muscular layer, cm, is of only moderate thickness and
is of rather a loose character. In the pyloric region it is not very
distinct from the underlying oblique layer, but in the other regions, as
has just been said, it is separated from the oblique layer by a
considerable layer of connective tissue like that of the submucosa.
The oblique layer, om, even in this section of the pyloric region is
the thickest of the three muscle layers; while in the cardiac, and
especially in the fundic, regions it is of great thickness, as was noted
above, and is made up of larger bundles with less intervening
connective tissue.
 Fig. 45. A transsection through the wall of the
pyloric region of the stomach of the feeding animal,
under low magnification; m, mucosa; om, oblique
muscles; other letters as in Figure 42.
 The outer or longitudinal muscle layer, lm, is comparatively little
developed and consists of small rather scattered bundles of muscles
with a correspondingly large amount of connective tissue. This
connective tissue passes insensibly into that of the surrounding
serosa, s, a loose, vascular layer of varying thickness and density,
shown very thick in Figure 45, but often much thinner.
So far as could be determined, the mucous membrane has the
same structure in both anterior and middle regions of the stomach.
That of the pyloric or small region, although fixed, stained, et cetera,
just as carefully as the rest, did not show cell details sufficiently well
to draw; the ducts of the glands in this region are fairly distinct but
the deeper parts of the glands have the appearance of series of
alveoli or large adipose cells. What the significance of this condition
may be the writer is not able to say, but since the structure of this
region of the gastric mucous membrane is not clear no attempt will
be made to describe its appearance under higher magnification than
was employed in the figure above. However, as will be noted below,
there is probably no great difference between the pyloric mucosa
and that of the other regions of the stomach.
 Fig. 46. The glands of the middle or fundic region of the stomach of the
hibernating animal, under high magnification; A, through duct; B, through body of
gland; C, through fundus of gland.
 Figure 46 shows portions of typical glands from the mucosa of the
middle region of the stomach, the posterior border of the large
stomach cavity; A is a longitudinal section through two ducts where
they open to the surface; B is a similar section through the body of a
gland below the region of the duct; C is a transsection through the
bottom or fundus of a gland; all are drawn with a camera under the
same magnification.
As is seen in Figure 45, under low magnification, the duct is about
one third of the entire length of the gland. The lumen of the duct is
fairly wide, that of the body of the gland is reduced to a mere slit,
while that of the fundus is quite wide.
One, two, or possibly more, glands may open to the surface
through one duct, as is shown in Figure 46. There is nothing peculiar
about the epithelium of these glands. Near the opening of the duct
the cells are of a typical columnar character with finely granular
cytoplasm, each with a nucleus at its basal end.
In the deeper parts of the duct the cells become shorter until in the
body of the gland (Fig. 46, B) they are cuboidal in outline.
The bodies of the glands are so closely packed together that it is
difficult to pick out an individual tube that will show details clearly
enough to draw with a camera lucida. So far as could be observed
all of the cells of this region of the gland are alike.
The bottom or fundus of the gland, as seen in Figure 46, C, is
somewhat enlarged and has a wide lumen. The cells are of the same
general character as in the more distal parts of the gland except that
they are somewhat more columnar or pyramidal than in the body of
the gland. The nuclei of the body and fundus are usually somewhat
larger and more nearly spherical than in the columnar cells of the
duct.
The feeding animals from which tissues were taken were
considerably smaller than the hibernating specimen, so that the
stomach walls were proportionately thinner; but, so far as could be
discovered, there was no difference in structure.
The relative thickness of the entire wall in each of the three
regions sectioned was about the same as described above.
As has been said, the mucosa on the pyloric or small region of the
stomach from the hibernating animal was so poorly fixed that its
structure could not be made out. In the feeding stage the mucosa of
this region was as well fixed as any of the other tissues and showed
that its structure is essentially like that shown in Figure 46, except
that the glands are proportionately not quite so long as in the fundic
and cardiac regions, and are somewhat more open—that is, they
have wider lumina; their lining cells are all of one kind and are
unchanged from what was seen in the hibernating condition.
The Small Intestine. Three regions of the small intestine will be
described: (1) an anterior, just caudad to the stomach; (2) a middle;
and (3) a posterior, one half inch cephalad to the rectum or large
intestine (Fig. 35).
As might be expected, the general structure of the wall of the
intestine is essentially the same in all three regions, the slight
differences noticeable being due mainly to variations in the thickness
of the various layers.
The middle and posterior regions have about the same diameter,
while the diameter of the anterior region is considerably greater, due
partly to the greater diameter of the lumen but mainly to the greater
thickness of the constituent layers, especially the mucosa. The
mucosa is also thrown into more numerous and complicated folds in
the anterior than in the middle and posterior regions; the complexity
of the mucosa seems to diminish as the intestine is followed caudad.
In the anterior region the mucosa may form at least one half of the
entire thickness of the wall, while in the posterior region it may form
less than one third of the thickness of the intestinal wall. The minute
structure of the intestinal epithelium will be described below.
The chief peculiarity of the intestinal wall is the apparent total
absence of a submucosa (Fig. 47). As will be described later, the
mucosal epithelium is laid upon the usual bed of fibrous and
lymphatic tissue, the tunica propria (Fig. 47, tp).
At the outer border of the tunica propria, and with no tissue
corresponding to a submucosa between it and the circular muscular
layer, is a thin and indistinct layer that has the appearance of a
longitudinal layer of muscle fibers; this should correspond to the
muscularis mucosa (Figs. 47, 48, 49, and 51, mm).
 Fig. 47. A transsection of the wall of the anterior region of the small intestine of
the hibernating animal, under low magnification; ln, lymph node; tp, tunica propria;
other letters as in Figure 42.

The circular, cm, and longitudinal, lm, muscle layers are compact,
and are distinct from the other layers of the wall; the former is
approximately twice the thickness of the latter. The relative thickness
of all the layers in the three regions of the intestine may be seen by
comparing Figures 47, 48, and 49.
 Fig. 48. An outline of a transsection of the wall of the middle
region of the small intestine of the hibernating animal, under low
magnification; lettering as in Figure 42.

The serosa, s, which is of about the same character in the three

regions under discussion, is a distinct and fairly dense layer of
connective tissue with numerous blood-vessels.
The general appearance of the mucous membrane as a whole is
sufficiently clear in the low-power drawing described above, so that
all that need be shown under a higher magnification is the epithelium
(Fig. 50). The upper part of this figure represents the lower end of
one of the intestinal glands cut longitudinally, below which is the end
of another gland in transverse section. Between the two sections is
the compact tunica propria of lymphatic tissue.
The section from which this particular figure was drawn was in the
anterior region, but the corresponding part of a section in either of
the other regions would have practically the same appearance.
The epithelium is of the stratified columnar type. The superficial
cells are very tall and narrow, with the nuclei generally at or near the
bases, though an occasional nucleus may be seen near the free end
of a cell. Below the tall columnar cells are four or five rows of nuclei
which represent smaller, irregular cells, though the cell walls could
not always be determined between the closely packed nuclei. No
goblet cells are to be seen at any place.

Fig. 49. An outline of a transsection through the

wall of the posterior region of the small intestine of the
hibernating animal, under low magnification; lettering
as in Figure 42.
 The relative diameters of the three regions of the small intestine in
the feeding condition are about the same as noted for the
hibernating stage; that is, the anterior region has the greatest
diameter and the other regions are smaller and have about the same
average diameter.
The most marked difference between the intestine during
hibernation and feeding is in the relative thickness of the mucosa
and muscular layers. As described for the hibernating stage, so in
the feeding stage, the mucosa is relatively the thickest in the anterior
regions and diminishes in thickness caudad; but while, in the
hibernating stage, it forms, in the anterior region, as much as half of
the entire thickness of the wall, in the feeding condition it forms, in
the same region, at least two thirds of the entire wall and in the
middle and posterior regions more than half of the wall.
 Fig. 50. Part of the mucous membrane of the anterior region of
the small intestine of the hibernating animal, under high
magnification. The upper part of the figure shows a part of a gland
cut longitudinally, the lower part of the figure shows another gland
cut transversely; e, epithelium; tp, tunica propria.
 The feeding animals being the smaller, the diameter of the
intestine was considerably less than in the hibernating stage; but the
actual thickness of the mucosa was practically the same, so that the
difference in diameter was due to the difference in the thickness of
the muscular and fibrous layers. It is therefore probable that the
differences noted above are due rather to the differences in the size
of the animals from which the tissues were taken than to the different
conditions of hibernation and feeding. The point to be noticed is that
the increase in the diameter of the intestine is due almost if not
entirely to an increase in thickness of the connective tissue and
muscle layers.
No difference in the complexity of the folds of the mucosa of the
two stages can be noticed.
The thickness of the fibro-muscular part of the wall of the intestine
varies considerably on different sides of the same region, but it
consists of the same layers in about the same relative amounts.
 Fig. 51. An outline of a transsection of the wall of
the middle region of the small intestine of the feeding
animal, under low magnification; m, mucosa; other
letters as in Figure 42.
 Figure 51 represents in outline the wall of the middle region of the
small intestine during feeding.
The epithelium is of the same thickness in the two stages, and the
only difference in its character that can be seen under a high
magnification is that, in the middle region at least, the nuclei are not
crowded so close together at the basal ends of the cells as in the
hibernating stage but are scattered more towards their free ends.