Etextbook 978 0077837280 Ecology Concepts and Applications 7th Edition

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Concepts and Applications 7th Edition

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Contents vii
8.2 Mate Choice and Resource Provisioning

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

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

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

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

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

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

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

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

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

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

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

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

survivorship curve.
pi- Dall sheep surviving their

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

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

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

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

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

Linda Bruslind Oregon State University Thomas O. Crist Miami University
Sherri L. Buerdsell West Virginia Northern Community College Peter Alpert University of Massachusetts—Amherst
Carrie E. Burdzinski Delta College (University Center, Michigan) Mark Pyron Ball State University
William Dew Nipissing University Mary Bremigan Michigan State University
Harry G. Deneer University of Saskatchewan
Phil Denette Delgado Community College Reviewers for the Fifth Edition
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Elyce Ervin University of Toledo Peter E. Busher Boston University
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Christina Gan Highline Community College James A. Fordyce University of Tennessee
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xxii Preface
Craig E. Williamson Miami University of Ohio Thomas W. Jurik Iowa State University
Jianguo (Jingle) Wu Arizona State University Karen L. Kandl University of New Orleans
Douglas Zook Boston University Robert Keys Cornerstone University
Mark E. Knauss Shorter College
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Kenneth M. Duke Brevard College Tom Robertson Portland Community College
Andy Dyer University of South Carolina Bernadette M. Roche Loyola College in Maryland
Ginny L. Eckert University of Alaska Tatiana Roth Coppin State College
J. Nicholas Ehringer Hillsborough Community College Neil Sabine Indiana University East
George F. Estabrook University of Michigan Seema Sanjay Jejurikar Bellevue Community College
Richard S. Feldman Marist College Timothy Savisky University of Pittsburgh
Charles A. Francis University of Nebraska—Lincoln Josh Schimel University of California—Santa Barbara
Carl Freeman Wayne State University Michael G. Scott Lincoln University
J. Phil Gibson Agnes Scott College Erik R. Scully Towson University
Robert R. Glesener Brevard College Michael J. Sebetich William Paterson University
Michael L. Golden Grossmont College Walter M. Shriner Mount Hood Community College
Paul Grecay Salisbury University John Skillman California State University—San Bernardino
Lana Hamilton Northeast State Tech Community College Jerry M. Skinner Keystone College
Brian Helmuth University of South Carolina Garriet W. Smith University of South Carolina—Aiken
James R. Hodgson Saint Norbert College Stacy Smith Lexington Community College
Jeremiah N. Jarrett Central Connecticut State University Joseph Stabile Iona College
Krish Jayachandran Florida International University Alan Stam Capital University
Mark Jonasson Crafton Hills College Alan Stiven University of North Carolina—Chapel Hill
Preface xxiii
Eric D. Storie Roanoke-Chowan Community College Fred E. Wasserman Boston University

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

the science of ecology.
W
hat is ecology? Ecology, the study of relation-
CHAPTER CONCEPTS ships between organisms and the environment,
has been a focus for human study for as long as
1.1 Ecologists study environmental we have existed as a species. Our survival has depended upon
relationships ranging from those how well we could observe variations in the environment and
of individual organisms to factors predict the responses of organisms to those variations. The
influencing global-scale processes. 2 earliest hunters and gatherers had to know the habits of their
Concept 1.1 Review 3 animal prey and where to find food plants. Later, agricultur-
ists had to be aware of variations in weather and soils and of
1.2 Ecologists design their studies based on how such variation might affect crops and livestock.
their research questions, the temporal Today, most of earth’s human population live in cities and
and spatial scale of their studies, and most of us have little direct contact with nature. More than ever
available research tools. 3 before, though, the future of our species depends on how well we
Concept 1.2 Review 8 understand the relationships between organisms and the environ-
ment. Our species is rapidly changing earth’s environment, yet
Investigating the Evidence 1: we do not fully understand the consequences of these changes.
The Scientific Method—Questions For instance, human activity has increased the quantity of nitro-
and Hypotheses 9 gen cycling through the biosphere, changed land cover across
Summary 10 the globe, and increased the atmospheric concentration of CO2.
Key Terms 10 Changes such as these threaten the diversity of life on earth and
Review Questions 10 may endanger our life support system. Because of the rapid pace
of environmental change at the dawn of the twenty-first century,
it is imperative that we continue as ardent students of ecology.
1
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

1.2 Sampling Ecological Research
1. How does the level of ecological organization an ecolo- LEARNING OUTCOMES
gist studies influence the questions he or she poses?
After studying this section you should be able to do the following:
2. While an ecologist may focus on a particular level of
ecological organization shown in figure 1.1, might other 1.5 Describe some emerging frontiers in ecology.
levels of organization be relevant, for example, does an 1.6 Explain how the use of stable isotopes has extended
ecologist studying factors limiting numbers in a popula- what it is possible to know about the ecology of warblers.
tion of zebras need to consider the influences of interac- 1.7 Compare the spatial and temporal scales addressed
tions with other species or the influences of food on the by the research of Robert MacArthur, Nalini
survival of individuals? Nadkarni, and Margaret Davis.
Ecologists design their studies based on their research warblers might be able to coexist and not compete with each
questions, the temporal and spatial scale of their studies, and other if they fed on the insects living in different zones within
available research tools. Because the discipline is so broad, trees. To map where the warblers fed, he subdivided trees into
ecological research can draw from all the physical and biologi- vertical and horizontal zones. He then carefully recorded the
cal sciences. The following section of this chapter provides a amount of time warblers spent feeding in each.
sample of ecological questions and approaches to research. 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
The Ecology of Forest Birds: figure 1.3 shows, the Cape May warbler fed mainly among
Old Tools and New new needles and buds at the tops of trees. The feeding zone
Robert MacArthur gazed intently through his binoculars. He of the blackburnian warbler overlapped broadly with that of
was watching a small bird, called a warbler, searching for the Cape May warbler but extended farther down the tree. The
insects in the top of a spruce tree. To the casual observer it might black-throated green warbler fed toward the trees’ interiors.
have seemed that MacArthur was a weekend bird-watcher. Yes, The bay-breasted warbler concentrated its feeding in the inte-
he was intensely interested in the birds he was watching, but he rior of trees. Finally, the yellow-rumped warbler fed mostly
was just as interested in testing ecological theory. on the ground and low in the trees. MacArthur’s observations
The year was 1955, and MacArthur was studying the showed that though these warblers live in the same forest,
ecology of five species of warblers that live together in the they extract food from different parts of that forest. He con-
spruce forests of northeastern North America. All five warbler cluded that feeding in different zones may reduce competition
species, Cape May (Dendroica tigrina), yellow-rumped among the warblers of spruce forests.
(D. coronata), black-throated green (D. virens), blackbur- MacArthur’s study (1958) of foraging by warblers is
nian (D. fusca), and bay-breasted (D. castanea), are about a true classic in the history of ecology. However, like most
the same size and shape and all feed on insects. Theory pre- studies it raised as many questions as it answered. Scientific
dicted that two species with identical ecological requirements research is important both for what it teaches us directly about
would compete with each other and that, as a consequence, nature and for how it stimulates other studies that improve
they could not live in the same environment indefinitely. Mac- our understanding. MacArthur’s work stimulated numerous
Arthur wanted to understand how several warbler species with studies of competition among many groups of organisms,
apparently similar ecological requirements could live together including warblers. Some of these studies produced results
in the same forest. that supported his work and others produced different results.
The warblers fed mainly by gleaning insects from the All added to our knowledge of competition between species
bark and foliage of trees. MacArthur predicted that these and of warbler ecology.
Cape May warbler Blackburnian warbler Black-throated

green warbler
New needles and New needles and buds New needles and buds
buds at top of tree of upper branches and some older needles
Bay-breasted Yellow-rumped
warbler warbler
Old needles and bare Bare or lichen-covered lower

and lichen-covered trunk and middle branches
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.
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.
Figure 1.4 A male American redstart, Setophaga ruticilla.

Mature male American redstarts are highly territorial, dominating Breeding grounds
high-quality feeding territories in their tropical wintering grounds,
pushing most female redstarts and young males into poorer-quality Wintering grounds
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 Figure 1.5 Map of the breeding and wintering grounds of the
(Norris et al. 2005) worked to develop tools capable of pen- American redstart, Setophaga ruticilla.
etrating the feeding habitats of wide-ranging migratory birds.
The object of their study was the American redstart (Setophaga Often, ecologists have pioneered the use of more power-
ruticilla), another colorful member of the warbler family Paru- ful research tools, as the complexity of their questions have
lidae (fig. 1.4). American redstarts, like the warblers studied by increased. A tool to which ecologists turn increasingly to
MacArthur, are long-distance migrants, nesting in temperate understand the ecology of migratory birds is stable isotope
North America but spending their winters mainly in tropical Cen- analysis (see chapter 6, p. 145). Isotopes of a chemical ele-
tral America, northern South America, and the Caribbean islands. ment, for example, isotopes of carbon, have different atomic
Historically, studies of wide-ranging bird species, such masses as a result of having different numbers of neutrons.
as the American redstart, have focused mainly on their tem- Carbon, for instance, has three isotopes (listed in order of
perate breeding grounds. However, observations by ecolo- increasing mass): 12C, 13C, and 14C. Of these three, 12C and
13
gists had long suggested that the success of an individual C are stable isotopes because they do not undergo radioac-
migratory bird during the breeding season may depend criti- tive decay, whereas 14C decays radioactively and is therefore
cally on the environmental conditions it experienced on its unstable. Stable isotopes have proven useful in the study of
tropical wintering grounds. For example, it has been well ecological processes—for example, identifying food sources,
established that male migratory birds, arriving early on the because the proportions of various isotopes differ across the
breeding grounds, are generally in better physical condition environment.
compared to those arriving later. Early arrivals also gen- Stable isotope analysis provides ecologists with a new
erally obtain the best breeding territories and have higher type of “lens” capable of revealing ecological relationships
reproductive success. that would otherwise remain invisible. For example, ecolo-
Variation in arrival times and physical condition led ecol- gists using stable isotope analysis can track habitat use by
ogists to ponder the connection between events on the win- American redstarts on their wintering grounds. In Jamaica,
tering grounds and subsequent reproductive success among older male American redstarts, along with some females,
birds in their breeding habitats. To answer such a question, spend the winter in higher-productivity mangrove forest hab-
we need a great deal of information, including where indi- itats, pushing most females and younger males into poorer-
vidual birds live on the wintering grounds, how the winter quality, dry scrub habitat. The dominant plants in these two
habitat correlates with physical condition during migration, habitats and the insects that feed on them contain different
how winter habitat influences time of arrival on the breeding proportions of the carbon isotopes 12C and 13C. Therefore,
grounds, and whether winter habitat correlates with reproduc- the tissues of the birds spending their winters in the produc-
tive success on the breeding grounds. Clearly, the amount of tive mangrove habitat (lower 13C) and those spending the
information required to answer such questions, concerning winters in the poor scrub habitat (higher 13C) are in effect
environments separated by thousands of kilometers (fig. 1.5), chemically tagged. As a consequence, today’s ecologist can
exceeds what one person, or even a large team, can learn analyze a very small sample of blood from an American red-
through the lenses of binoculars. start when it arrives on its temperate breeding ground and
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 Figure 1.6 Exploring the rain forest canopy. What Nalini Nadkarni
of how tropical and temperate rain forests are structured discovered helped solve an ecological puzzle.
and how they function. With the aid of mountain-climbing
equipment, Nadkarni slowly made her first ascent into the Epiphyte mats contain significant quantities of nutrients.
canopy of the Costa Rican rain forest, a world explored by Nadkarni estimated that these quantities in some tropical rain
few others and where she was to become a pioneer (fig. 1.6). forests are equal to about half the nutrient content of the foli-
She stood on the rain forest floor and wondered about the age of the canopy trees. In the temperate rain forests of the
diversity of organisms and ecological relationships that Olympic Peninsula in Washington, the mass of epiphytes is
might be hidden in the canopy high above. Her wonder soon four times the mass of leaves on their host trees.
gave way to determination, and Nadkarni not only visited Nadkarni’s research showed that in both temperate and
the canopy but was among the first to explore the ecology of tropical rain forests, trees access these nutrient stores by send-
this unseen world. ing out roots from their trunks and branches high above the
Because of leaching by heavy rains, many rain forest ground. These roots grow into the epiphyte mats and extract
soils are poor in nutrients such as nitrogen and phosphorus. nutrients from them. As a consequence of this research, we
The low availability of nutrients in many rain forest soils has now know that to understand the nutrient economy of rain for-
produced one of ecology’s puzzles. How can the prodigious ests the ecologist must venture into the treetops.
life of rain forests be maintained on such nutrient-poor soils? Easier means of working in the rain forest canopy have
Many factors contribute to the maintenance of this intense been developed, and this research is no longer limited to the
biological activity. Nadkarni’s research in the treetops uncov- adventurous and agile. New ways to access the forest canopy
ered one of those factors, a significant store of nutrients in the range from hot air balloons and aerial trams to large cranes.
rain forest canopy. The Wind River Canopy Crane offers scientists access to any
The nutrient stores in the rain forest canopy are associ- level within a 70 m tall coniferous forest in a 2.3 ha area near
ated with epiphytes. Epiphytes are plants, such as many the Columbia River Gorge in Washington (fig. 1.7). Research
orchids and ferns, that live on the branches and trunks of other projects supported—and made far easier—by this crane and
plants. Epiphytes are not parasitic: they do not derive their others have included the ecology of migratory birds in the
nutrition from the plant they grow on. As they grow on the forest canopy, photosynthesis by epiphytes living at differ-
branches of a tree they begin to trap organic matter, which ent canopy heights, and vertical stratification of habitat use
eventually forms a mat. Epiphyte mats increase in thickness by bats and beetles (Ozanne et al. 2003). By 2006, there were
up to 30 cm, providing a complex structure that supports a 12 canopy cranes facilitating canopy research in temperate
diverse community of plants and animals. and tropical forests worldwide (Stork 2007). Nadkarni points
Gondola housing scientists can Crane operator can swing crane

Canopy zonation be lowered to study any level in arm 360˚ and position gondola
the canopy. at any distance along crane arm.
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 Pacific Grand Pacific Western Pacific Western
American robin, winter wren, black- fir silver fir fir yew hemlock dogwood red cedar
tailed deer, coyote
Tree species
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 lake sediments. As lake sediments build up over the centu-
physical frontier may be closing, but its exploration as a sci- ries, this pollen is preserved and forms a historical record
entific frontier is just beginning, particularly as we attempt to of the kinds of plants that lived nearby. As the lakeside veg-
predict the ecological consequences of climate change. 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
Climatic and Ecological Change: sediments about 12,000 years ago then pollen from beech,
Past and Future Fagus grandifolia, occurs in the sediments beginning
The earth and its life are always changing. However, many about 8,000 years ago. Chestnut pollen does not appear
of the most important changes occur over such long periods in the sediments until about 2,000 years ago. The pollen
of time or at such large spatial scales that they are difficult from all three tree species continues in the sediment record
to study. Two approaches that provide insights into long-term until about 1920, when chestnut blight killed most of the
and large-scale processes are studies of pollen preserved in chestnut trees in the vicinity of the lake. Thus, the pollen
lake sediments and evolutionary studies. preserved in the sediments of lakes can be used to recon-
Margaret Davis (1983, 1989) carefully searched through struct the history of vegetation in the area. Margaret B.
a sample of lake sediments for pollen. The sediments had Davis, Ruth G. Shaw, and Julie R. Etterson review extensive
come from a lake in the Appalachian Mountains, and the evidence that during climate change, plants evolve, as well
pollen they contained would help her document changes in as disperse (Davis and Shaw 2001; Davis, Shaw, and Etter-
the community of plants living near the lake during the past son 2005). As climate changes, plant populations simultane-
several thousand years. Davis is a paleoecologist trained to ously change their geographic distributions and undergo the
think at very large spatial scales and over very long periods of evolutionary process of adaptation, which increases their
time. She has spent much of her professional career studying ability to live in the new climatic regime. Meanwhile, evi-
changes in the distributions of plants during the Quaternary dence of evolutionary responses to climate change is being
period, particularly during the most recent 20,000 years. discovered among many animal groups. William Bradshaw
Some of the pollen produced by plants that live near a and Christina Holzapfel (2006) summarized several stud-
lake falls on the lake surface, sinks, and becomes trapped in ies documenting evolutionary change in northern animals,
Lake profile Chestnut decline Present Pollen

due to blight 100
Chestnut
2,000
Years
Beech
8,000
12,000 Spruce
Sediments
Sediment core
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), warming (see chapter 23, p. 519). Research such as that
in response to increasing growing season length as a conse- by Davis and her colleagues will be essential to predicting
quence of the now-well-documented phenomenon of global 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
Figure 1.9 Studies indicate that north American red squirrels, her colleagues did not identify the questions that they
Tamiasciurus hudsonicus, have been undergoing rapid evolution for addressed. What research questions can we infer from
earlier breeding, during a recent period of increased average spring the above description of their work?
temperatures in Canada’s Yukon Territory (Réale et al. 2003).
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
A hypothesis is a possible answer to a question. MacArthur’s
main hypothesis (possible answer to his question) was: “Sev-
Questions and Hypotheses eral warbler species are able to coexist because each species
What do scientists do? Simply put, scientists ask and attempt feeds on insects living in different zones within trees.”
to find answers to questions about the natural world. Ques- Once a scientist or team of scientists proposes a hypoth-
tions are the guiding lights of the scientific process. With- esis (or multiple alternative hypotheses), the next step in the
out them, exploration of nature lacks focus and yields little scientific method is to determine its validity by testing predic-
understanding of the world. Let’s consider a question asked tions that follow from the hypothesis. Three fundamental ways
by an ecologist discussed in this chapter. The main ques- to test hypotheses are through observation, experiments, and
tion asked by Robert MacArthur in his studies of warblers modeling. These approaches, which are all represented in fig-
(p. 4) was something like the following: “How can several ure 1, will be discussed in detail in the “Investigating the Evi-
species of insect-eating warblers live in the same forest with- dence” boxes and in the research discussed in later chapters.
out one species eventually excluding the others through com-
petition?” While this focus on questions may seem obvious, CRITIQUING THE EVIDENCE 1
one of the most common questions asked of scientists at semi- 1. How does the development of new research tools, such as
nars and professional meetings is, “What is your question?” canopy cranes and stable isotope analysis, affect the pro-
If scientists are in the business of asking questions cess of science as outlined by figure 1 of this “Investigat-
about nature, where does a hypothesis enter the process? ing the Evidence” box?
Summary
Ecologists study environmental relationships ranging from reviewed, ecologists study those relationships over a large
those of individual organisms to factors influencing global- range of temporal and spatial scales using a wide variety of
scale processes. The research focus and questions posed by approaches. Ecology includes Davis’s studies of vegetation
ecologists differ across the levels of ecological organization moving across the North American continent over a span of
studied. thousands of years. Ecology also includes the observational
Ecologists design their studies based on their research studies of birds in contemporary forests by MacArthur. Ecolo-
questions, the temporal and spatial scale of their stud- gists may study processes on plots measured in square centime-
ies, and available research tools. With this brief review ters or, like those studying the ecology of migratory birds, study
of research approaches and topics, we return to the question areas may span thousands of kilometers. Important ecological
asked at the beginning of the chapter: What is ecology? Ecol- discoveries have come from Nadkarni’s probing of the rain for-
ogy is indeed the study of relationships between organisms est canopy and from traces of stable isotopes in a droplet of
and the environment. However, as you can see from the studies blood. Ecology includes all these approaches and many more.
Key Terms
adaptation 7 ecology 1 evolution 2 stable isotope analysis 5
aeroecology 3 ecosystem 6 nutrient 6 urban ecology 3
biosphere 3 epiphyte 6
Review Questions
1. Faced with the complexity of nature, ecologists have divided the of nutrient storage in rain forest canopy resulted from the biol-
field of ecology into subdisciplines, each of which focuses on ogy of individual organisms, populations of organisms, and
one of the levels of organization pictured in figure 1.1. What is communities of species. Explain.
the advantage of developing such subdisciplines within ecology? 5. What do the studies of Margaret Davis tell us about the compo-
2. What are the pitfalls of subdividing nature in the way it is repre- sition of forests in the Appalachian Mountains during the past
sented in figure 1.1? In what ways does figure 1.1 misrepresent 12,000 years (see fig. 1.8)? Based on this research, what pre-
nature? dictions might you make about the future composition of these
3. What could you do to verify that the distinct feeding zones forests?
used by the warblers studied by MacArthur (see fig. 1.3) are 6. During the course of the studies reviewed in this chapter, each
the result of ongoing competition between the different species scientist or team of scientists measured certain variables. What
of warblers? How might you examine the role of competition major variable studied by Margaret Davis and her research
in keeping some American redstarts out of the most productive team distinguishes their work from that of the other research
feeding areas on their wintering grounds? reviewed in the chapter?
4. Although Nalini Nadkarni’s studies of the rain forest canopy
addressed a question related to ecosystem structure, the patterns
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DANCE ON STILTS AT THE GIRLS’ UNYAGO, NIUCHI
Newala, too, suffers from the distance of its water-supply—at least

the Newala of to-day does; there was once another Newala in a lovely
valley at the foot of the plateau. I visited it and found scarcely a trace
of houses, only a Christian cemetery, with the graves of several
missionaries and their converts, remaining as a monument of its
former glories. But the surroundings are wonderfully beautiful. A
thick grove of splendid mango-trees closes in the weather-worn
crosses and headstones; behind them, combining the useful and the
agreeable, is a whole plantation of lemon-trees covered with ripe
fruit; not the small African kind, but a much larger and also juicier
imported variety, which drops into the hands of the passing traveller,
without calling for any exertion on his part. Old Newala is now under
the jurisdiction of the native pastor, Daudi, at Chingulungulu, who,
as I am on very friendly terms with him, allows me, as a matter of
course, the use of this lemon-grove during my stay at Newala.
FEET MUTILATED BY THE RAVAGES OF THE “JIGGER”
(Sarcopsylla penetrans)
The water-supply of New Newala is in the bottom of the valley,

some 1,600 feet lower down. The way is not only long and fatiguing,
but the water, when we get it, is thoroughly bad. We are suffering not
only from this, but from the fact that the arrangements at Newala are
nothing short of luxurious. We have a separate kitchen—a hut built
against the boma palisade on the right of the baraza, the interior of
which is not visible from our usual position. Our two cooks were not
long in finding this out, and they consequently do—or rather neglect
to do—what they please. In any case they do not seem to be very
particular about the boiling of our drinking-water—at least I can
attribute to no other cause certain attacks of a dysenteric nature,
from which both Knudsen and I have suffered for some time. If a
man like Omari has to be left unwatched for a moment, he is capable
of anything. Besides this complaint, we are inconvenienced by the
state of our nails, which have become as hard as glass, and crack on
the slightest provocation, and I have the additional infliction of
pimples all over me. As if all this were not enough, we have also, for
the last week been waging war against the jigger, who has found his
Eldorado in the hot sand of the Makonde plateau. Our men are seen
all day long—whenever their chronic colds and the dysentery likewise
raging among them permit—occupied in removing this scourge of
Africa from their feet and trying to prevent the disastrous
consequences of its presence. It is quite common to see natives of
this place with one or two toes missing; many have lost all their toes,
or even the whole front part of the foot, so that a well-formed leg
ends in a shapeless stump. These ravages are caused by the female of
Sarcopsylla penetrans, which bores its way under the skin and there
develops an egg-sac the size of a pea. In all books on the subject, it is
stated that one’s attention is called to the presence of this parasite by
an intolerable itching. This agrees very well with my experience, so
far as the softer parts of the sole, the spaces between and under the
toes, and the side of the foot are concerned, but if the creature
penetrates through the harder parts of the heel or ball of the foot, it
may escape even the most careful search till it has reached maturity.
Then there is no time to be lost, if the horrible ulceration, of which
we see cases by the dozen every day, is to be prevented. It is much
easier, by the way, to discover the insect on the white skin of a
European than on that of a native, on which the dark speck scarcely
shows. The four or five jiggers which, in spite of the fact that I
constantly wore high laced boots, chose my feet to settle in, were
taken out for me by the all-accomplished Knudsen, after which I
thought it advisable to wash out the cavities with corrosive
sublimate. The natives have a different sort of disinfectant—they fill
the hole with scraped roots. In a tiny Makua village on the slope of
the plateau south of Newala, we saw an old woman who had filled all
the spaces under her toe-nails with powdered roots by way of
prophylactic treatment. What will be the result, if any, who can say?
The rest of the many trifling ills which trouble our existence are
really more comic than serious. In the absence of anything else to
smoke, Knudsen and I at last opened a box of cigars procured from
the Indian store-keeper at Lindi, and tried them, with the most
distressing results. Whether they contain opium or some other
narcotic, neither of us can say, but after the tenth puff we were both
“off,” three-quarters stupefied and unspeakably wretched. Slowly we
recovered—and what happened next? Half-an-hour later we were
once more smoking these poisonous concoctions—so insatiable is the
craving for tobacco in the tropics.
Even my present attacks of fever scarcely deserve to be taken
seriously. I have had no less than three here at Newala, all of which
have run their course in an incredibly short time. In the early
afternoon, I am busy with my old natives, asking questions and
making notes. The strong midday coffee has stimulated my spirits to
an extraordinary degree, the brain is active and vigorous, and work
progresses rapidly, while a pleasant warmth pervades the whole
body. Suddenly this gives place to a violent chill, forcing me to put on
my overcoat, though it is only half-past three and the afternoon sun
is at its hottest. Now the brain no longer works with such acuteness
and logical precision; more especially does it fail me in trying to
establish the syntax of the difficult Makua language on which I have
ventured, as if I had not enough to do without it. Under the
circumstances it seems advisable to take my temperature, and I do
so, to save trouble, without leaving my seat, and while going on with
my work. On examination, I find it to be 101·48°. My tutors are
abruptly dismissed and my bed set up in the baraza; a few minutes
later I am in it and treating myself internally with hot water and
lemon-juice.
Three hours later, the thermometer marks nearly 104°, and I make
them carry me back into the tent, bed and all, as I am now perspiring
heavily, and exposure to the cold wind just beginning to blow might
mean a fatal chill. I lie still for a little while, and then find, to my
great relief, that the temperature is not rising, but rather falling. This
is about 7.30 p.m. At 8 p.m. I find, to my unbounded astonishment,
that it has fallen below 98·6°, and I feel perfectly well. I read for an
hour or two, and could very well enjoy a smoke, if I had the
wherewithal—Indian cigars being out of the question.
Having no medical training, I am at a loss to account for this state
of things. It is impossible that these transitory attacks of high fever
should be malarial; it seems more probable that they are due to a
kind of sunstroke. On consulting my note-book, I become more and
more inclined to think this is the case, for these attacks regularly
follow extreme fatigue and long exposure to strong sunshine. They at
least have the advantage of being only short interruptions to my
work, as on the following morning I am always quite fresh and fit.
My treasure of a cook is suffering from an enormous hydrocele which
makes it difficult for him to get up, and Moritz is obliged to keep in
the dark on account of his inflamed eyes. Knudsen’s cook, a raw boy
from somewhere in the bush, knows still less of cooking than Omari;
consequently Nils Knudsen himself has been promoted to the vacant
post. Finding that we had come to the end of our supplies, he began
by sending to Chingulungulu for the four sucking-pigs which we had
bought from Matola and temporarily left in his charge; and when
they came up, neatly packed in a large crate, he callously slaughtered
the biggest of them. The first joint we were thoughtless enough to
entrust for roasting to Knudsen’s mshenzi cook, and it was
consequently uneatable; but we made the rest of the animal into a
jelly which we ate with great relish after weeks of underfeeding,
consuming incredible helpings of it at both midday and evening
meals. The only drawback is a certain want of variety in the tinned
vegetables. Dr. Jäger, to whom the Geographical Commission
entrusted the provisioning of the expeditions—mine as well as his
own—because he had more time on his hands than the rest of us,
seems to have laid in a huge stock of Teltow turnips,[46] an article of
food which is all very well for occasional use, but which quickly palls
when set before one every day; and we seem to have no other tins
left. There is no help for it—we must put up with the turnips; but I
am certain that, once I am home again, I shall not touch them for ten
years to come.
Amid all these minor evils, which, after all, go to make up the
genuine flavour of Africa, there is at least one cheering touch:
Knudsen has, with the dexterity of a skilled mechanic, repaired my 9
× 12 cm. camera, at least so far that I can use it with a little care.
How, in the absence of finger-nails, he was able to accomplish such a
ticklish piece of work, having no tool but a clumsy screw-driver for
taking to pieces and putting together again the complicated
mechanism of the instantaneous shutter, is still a mystery to me; but
he did it successfully. The loss of his finger-nails shows him in a light
contrasting curiously enough with the intelligence evinced by the
above operation; though, after all, it is scarcely surprising after his
ten years’ residence in the bush. One day, at Lindi, he had occasion
to wash a dog, which must have been in need of very thorough
cleansing, for the bottle handed to our friend for the purpose had an
extremely strong smell. Having performed his task in the most
conscientious manner, he perceived with some surprise that the dog
did not appear much the better for it, and was further surprised by
finding his own nails ulcerating away in the course of the next few
days. “How was I to know that carbolic acid has to be diluted?” he
mutters indignantly, from time to time, with a troubled gaze at his
mutilated finger-tips.
Since we came to Newala we have been making excursions in all
directions through the surrounding country, in accordance with old
habit, and also because the akida Sefu did not get together the tribal
elders from whom I wanted information so speedily as he had
promised. There is, however, no harm done, as, even if seen only
from the outside, the country and people are interesting enough.
The Makonde plateau is like a large rectangular table rounded off
at the corners. Measured from the Indian Ocean to Newala, it is
about seventy-five miles long, and between the Rovuma and the
Lukuledi it averages fifty miles in breadth, so that its superficial area
is about two-thirds of that of the kingdom of Saxony. The surface,
however, is not level, but uniformly inclined from its south-western
edge to the ocean. From the upper edge, on which Newala lies, the
eye ranges for many miles east and north-east, without encountering
any obstacle, over the Makonde bush. It is a green sea, from which
here and there thick clouds of smoke rise, to show that it, too, is
inhabited by men who carry on their tillage like so many other
primitive peoples, by cutting down and burning the bush, and
manuring with the ashes. Even in the radiant light of a tropical day
such a fire is a grand sight.
Much less effective is the impression produced just now by the
great western plain as seen from the edge of the plateau. As often as
time permits, I stroll along this edge, sometimes in one direction,
sometimes in another, in the hope of finding the air clear enough to
let me enjoy the view; but I have always been disappointed.
Wherever one looks, clouds of smoke rise from the burning bush,
and the air is full of smoke and vapour. It is a pity, for under more
favourable circumstances the panorama of the whole country up to
the distant Majeje hills must be truly magnificent. It is of little use
taking photographs now, and an outline sketch gives a very poor idea
of the scenery. In one of these excursions I went out of my way to
make a personal attempt on the Makonde bush. The present edge of
the plateau is the result of a far-reaching process of destruction
through erosion and denudation. The Makonde strata are
everywhere cut into by ravines, which, though short, are hundreds of
yards in depth. In consequence of the loose stratification of these
beds, not only are the walls of these ravines nearly vertical, but their
upper end is closed by an equally steep escarpment, so that the
western edge of the Makonde plateau is hemmed in by a series of
deep, basin-like valleys. In order to get from one side of such a ravine
to the other, I cut my way through the bush with a dozen of my men.
It was a very open part, with more grass than scrub, but even so the
short stretch of less than two hundred yards was very hard work; at
the end of it the men’s calicoes were in rags and they themselves
bleeding from hundreds of scratches, while even our strong khaki
suits had not escaped scatheless.
NATIVE PATH THROUGH THE MAKONDE BUSH, NEAR

MAHUTA
I see increasing reason to believe that the view formed some time
back as to the origin of the Makonde bush is the correct one. I have
no doubt that it is not a natural product, but the result of human
occupation. Those parts of the high country where man—as a very
slight amount of practice enables the eye to perceive at once—has not
yet penetrated with axe and hoe, are still occupied by a splendid
timber forest quite able to sustain a comparison with our mixed
forests in Germany. But wherever man has once built his hut or tilled
his field, this horrible bush springs up. Every phase of this process
may be seen in the course of a couple of hours’ walk along the main
road. From the bush to right or left, one hears the sound of the axe—
not from one spot only, but from several directions at once. A few
steps further on, we can see what is taking place. The brush has been
cut down and piled up in heaps to the height of a yard or more,
between which the trunks of the large trees stand up like the last
pillars of a magnificent ruined building. These, too, present a
melancholy spectacle: the destructive Makonde have ringed them—
cut a broad strip of bark all round to ensure their dying off—and also
piled up pyramids of brush round them. Father and son, mother and
son-in-law, are chopping away perseveringly in the background—too
busy, almost, to look round at the white stranger, who usually excites
so much interest. If you pass by the same place a week later, the piles
of brushwood have disappeared and a thick layer of ashes has taken
the place of the green forest. The large trees stretch their
smouldering trunks and branches in dumb accusation to heaven—if
they have not already fallen and been more or less reduced to ashes,
perhaps only showing as a white stripe on the dark ground.
This work of destruction is carried out by the Makonde alike on the
virgin forest and on the bush which has sprung up on sites already
cultivated and deserted. In the second case they are saved the trouble
of burning the large trees, these being entirely absent in the
secondary bush.
After burning this piece of forest ground and loosening it with the
hoe, the native sows his corn and plants his vegetables. All over the
country, he goes in for bed-culture, which requires, and, in fact,
receives, the most careful attention. Weeds are nowhere tolerated in
the south of German East Africa. The crops may fail on the plains,
where droughts are frequent, but never on the plateau with its
abundant rains and heavy dews. Its fortunate inhabitants even have
the satisfaction of seeing the proud Wayao and Wamakua working
for them as labourers, driven by hunger to serve where they were
accustomed to rule.
But the light, sandy soil is soon exhausted, and would yield no
harvest the second year if cultivated twice running. This fact has
been familiar to the native for ages; consequently he provides in
time, and, while his crop is growing, prepares the next plot with axe
and firebrand. Next year he plants this with his various crops and
lets the first piece lie fallow. For a short time it remains waste and
desolate; then nature steps in to repair the destruction wrought by
man; a thousand new growths spring out of the exhausted soil, and
even the old stumps put forth fresh shoots. Next year the new growth
is up to one’s knees, and in a few years more it is that terrible,
impenetrable bush, which maintains its position till the black
occupier of the land has made the round of all the available sites and
come back to his starting point.
The Makonde are, body and soul, so to speak, one with this bush.
According to my Yao informants, indeed, their name means nothing
else but “bush people.” Their own tradition says that they have been
settled up here for a very long time, but to my surprise they laid great
stress on an original immigration. Their old homes were in the
south-east, near Mikindani and the mouth of the Rovuma, whence
their peaceful forefathers were driven by the continual raids of the
Sakalavas from Madagascar and the warlike Shirazis[47] of the coast,
to take refuge on the almost inaccessible plateau. I have studied
African ethnology for twenty years, but the fact that changes of
population in this apparently quiet and peaceable corner of the earth
could have been occasioned by outside enterprises taking place on
the high seas, was completely new to me. It is, no doubt, however,
correct.
The charming tribal legend of the Makonde—besides informing us
of other interesting matters—explains why they have to live in the
thickest of the bush and a long way from the edge of the plateau,
instead of making their permanent homes beside the purling brooks
and springs of the low country.
“The place where the tribe originated is Mahuta, on the southern
side of the plateau towards the Rovuma, where of old time there was
nothing but thick bush. Out of this bush came a man who never
washed himself or shaved his head, and who ate and drank but little.
He went out and made a human figure from the wood of a tree
growing in the open country, which he took home to his abode in the
bush and there set it upright. In the night this image came to life and
was a woman. The man and woman went down together to the
Rovuma to wash themselves. Here the woman gave birth to a still-
born child. They left that place and passed over the high land into the
valley of the Mbemkuru, where the woman had another child, which
was also born dead. Then they returned to the high bush country of
Mahuta, where the third child was born, which lived and grew up. In
course of time, the couple had many more children, and called
themselves Wamatanda. These were the ancestral stock of the
Makonde, also called Wamakonde,[48] i.e., aborigines. Their
forefather, the man from the bush, gave his children the command to
bury their dead upright, in memory of the mother of their race who
was cut out of wood and awoke to life when standing upright. He also
warned them against settling in the valleys and near large streams,
for sickness and death dwelt there. They were to make it a rule to
have their huts at least an hour’s walk from the nearest watering-
place; then their children would thrive and escape illness.”
The explanation of the name Makonde given by my informants is
somewhat different from that contained in the above legend, which I
extract from a little book (small, but packed with information), by
Pater Adams, entitled Lindi und sein Hinterland. Otherwise, my
results agree exactly with the statements of the legend. Washing?
Hapana—there is no such thing. Why should they do so? As it is, the
supply of water scarcely suffices for cooking and drinking; other
people do not wash, so why should the Makonde distinguish himself
by such needless eccentricity? As for shaving the head, the short,
woolly crop scarcely needs it,[49] so the second ancestral precept is
likewise easy enough to follow. Beyond this, however, there is
nothing ridiculous in the ancestor’s advice. I have obtained from
various local artists a fairly large number of figures carved in wood,
ranging from fifteen to twenty-three inches in height, and
representing women belonging to the great group of the Mavia,
Makonde, and Matambwe tribes. The carving is remarkably well
done and renders the female type with great accuracy, especially the
keloid ornamentation, to be described later on. As to the object and
meaning of their works the sculptors either could or (more probably)
would tell me nothing, and I was forced to content myself with the
scanty information vouchsafed by one man, who said that the figures
were merely intended to represent the nembo—the artificial
deformations of pelele, ear-discs, and keloids. The legend recorded
by Pater Adams places these figures in a new light. They must surely
be more than mere dolls; and we may even venture to assume that
they are—though the majority of present-day Makonde are probably
unaware of the fact—representations of the tribal ancestress.
The references in the legend to the descent from Mahuta to the
Rovuma, and to a journey across the highlands into the Mbekuru
valley, undoubtedly indicate the previous history of the tribe, the
travels of the ancestral pair typifying the migrations of their
descendants. The descent to the neighbouring Rovuma valley, with
its extraordinary fertility and great abundance of game, is intelligible
at a glance—but the crossing of the Lukuledi depression, the ascent
to the Rondo Plateau and the descent to the Mbemkuru, also lie
within the bounds of probability, for all these districts have exactly
the same character as the extreme south. Now, however, comes a
point of especial interest for our bacteriological age. The primitive
Makonde did not enjoy their lives in the marshy river-valleys.
Disease raged among them, and many died. It was only after they
had returned to their original home near Mahuta, that the health
conditions of these people improved. We are very apt to think of the
African as a stupid person whose ignorance of nature is only equalled
by his fear of it, and who looks on all mishaps as caused by evil
spirits and malignant natural powers. It is much more correct to
assume in this case that the people very early learnt to distinguish
districts infested with malaria from those where it is absent.
This knowledge is crystallized in the
ancestral warning against settling in the
valleys and near the great waters, the
dwelling-places of disease and death. At the
same time, for security against the hostile
Mavia south of the Rovuma, it was enacted
that every settlement must be not less than a
certain distance from the southern edge of the
plateau. Such in fact is their mode of life at the
present day. It is not such a bad one, and
certainly they are both safer and more
comfortable than the Makua, the recent
intruders from the south, who have made USUAL METHOD OF
good their footing on the western edge of the CLOSING HUT-DOOR
plateau, extending over a fairly wide belt of
country. Neither Makua nor Makonde show in their dwellings
anything of the size and comeliness of the Yao houses in the plain,
especially at Masasi, Chingulungulu and Zuza’s. Jumbe Chauro, a
Makonde hamlet not far from Newala, on the road to Mahuta, is the
most important settlement of the tribe I have yet seen, and has fairly
spacious huts. But how slovenly is their construction compared with
the palatial residences of the elephant-hunters living in the plain.
The roofs are still more untidy than in the general run of huts during
the dry season, the walls show here and there the scanty beginnings
or the lamentable remains of the mud plastering, and the interior is a
veritable dog-kennel; dirt, dust and disorder everywhere. A few huts
only show any attempt at division into rooms, and this consists
merely of very roughly-made bamboo partitions. In one point alone
have I noticed any indication of progress—in the method of fastening
the door. Houses all over the south are secured in a simple but
ingenious manner. The door consists of a set of stout pieces of wood
or bamboo, tied with bark-string to two cross-pieces, and moving in
two grooves round one of the door-posts, so as to open inwards. If
the owner wishes to leave home, he takes two logs as thick as a man’s
upper arm and about a yard long. One of these is placed obliquely
against the middle of the door from the inside, so as to form an angle
of from 60° to 75° with the ground. He then places the second piece
horizontally across the first, pressing it downward with all his might.
It is kept in place by two strong posts planted in the ground a few
inches inside the door. This fastening is absolutely safe, but of course
cannot be applied to both doors at once, otherwise how could the
owner leave or enter his house? I have not yet succeeded in finding
out how the back door is fastened.
MAKONDE LOCK AND KEY AT JUMBE CHAURO

This is the general way of closing a house. The Makonde at Jumbe
Chauro, however, have a much more complicated, solid and original
one. Here, too, the door is as already described, except that there is
only one post on the inside, standing by itself about six inches from
one side of the doorway. Opposite this post is a hole in the wall just
large enough to admit a man’s arm. The door is closed inside by a
large wooden bolt passing through a hole in this post and pressing
with its free end against the door. The other end has three holes into
which fit three pegs running in vertical grooves inside the post. The
door is opened with a wooden key about a foot long, somewhat
curved and sloped off at the butt; the other end has three pegs
corresponding to the holes, in the bolt, so that, when it is thrust
through the hole in the wall and inserted into the rectangular
opening in the post, the pegs can be lifted and the bolt drawn out.[50]
MODE OF INSERTING THE KEY
With no small pride first one householder and then a second

showed me on the spot the action of this greatest invention of the
Makonde Highlands. To both with an admiring exclamation of
“Vizuri sana!” (“Very fine!”). I expressed the wish to take back these
marvels with me to Ulaya, to show the Wazungu what clever fellows
the Makonde are. Scarcely five minutes after my return to camp at
Newala, the two men came up sweating under the weight of two
heavy logs which they laid down at my feet, handing over at the same
time the keys of the fallen fortress. Arguing, logically enough, that if
the key was wanted, the lock would be wanted with it, they had taken
their axes and chopped down the posts—as it never occurred to them
to dig them out of the ground and so bring them intact. Thus I have
two badly damaged specimens, and the owners, instead of praise,
come in for a blowing-up.
The Makua huts in the environs of Newala are especially
miserable; their more than slovenly construction reminds one of the
temporary erections of the Makua at Hatia’s, though the people here
have not been concerned in a war. It must therefore be due to
congenital idleness, or else to the absence of a powerful chief. Even
the baraza at Mlipa’s, a short hour’s walk south-east of Newala,
shares in this general neglect. While public buildings in this country
are usually looked after more or less carefully, this is in evident
danger of being blown over by the first strong easterly gale. The only
attractive object in this whole district is the grave of the late chief
Mlipa. I visited it in the morning, while the sun was still trying with
partial success to break through the rolling mists, and the circular
grove of tall euphorbias, which, with a broken pot, is all that marks
the old king’s resting-place, impressed one with a touch of pathos.
Even my very materially-minded carriers seemed to feel something
of the sort, for instead of their usual ribald songs, they chanted
solemnly, as we marched on through the dense green of the Makonde
bush:—
“We shall arrive with the great master; we stand in a row and have
no fear about getting our food and our money from the Serkali (the
Government). We are not afraid; we are going along with the great
master, the lion; we are going down to the coast and back.”
With regard to the characteristic features of the various tribes here
on the western edge of the plateau, I can arrive at no other
conclusion than the one already come to in the plain, viz., that it is
impossible for anyone but a trained anthropologist to assign any
given individual at once to his proper tribe. In fact, I think that even
an anthropological specialist, after the most careful examination,
might find it a difficult task to decide. The whole congeries of peoples
collected in the region bounded on the west by the great Central
African rift, Tanganyika and Nyasa, and on the east by the Indian
Ocean, are closely related to each other—some of their languages are
only distinguished from one another as dialects of the same speech,
and no doubt all the tribes present the same shape of skull and
structure of skeleton. Thus, surely, there can be no very striking
differences in outward appearance.
Even did such exist, I should have no time
to concern myself with them, for day after day,
I have to see or hear, as the case may be—in
any case to grasp and record—an
extraordinary number of ethnographic
phenomena. I am almost disposed to think it
fortunate that some departments of inquiry, at
least, are barred by external circumstances.
Chief among these is the subject of iron-
working. We are apt to think of Africa as a
country where iron ore is everywhere, so to
speak, to be picked up by the roadside, and
where it would be quite surprising if the
inhabitants had not learnt to smelt the
material ready to their hand. In fact, the
knowledge of this art ranges all over the
continent, from the Kabyles in the north to the
Kafirs in the south. Here between the Rovuma
and the Lukuledi the conditions are not so
favourable. According to the statements of the
Makonde, neither ironstone nor any other
form of iron ore is known to them. They have
not therefore advanced to the art of smelting
the metal, but have hitherto bought all their
THE ANCESTRESS OF
THE MAKONDE
iron implements from neighbouring tribes.
Even in the plain the inhabitants are not much
better off. Only one man now living is said to
understand the art of smelting iron. This old fundi lives close to
Huwe, that isolated, steep-sided block of granite which rises out of
the green solitude between Masasi and Chingulungulu, and whose
jagged and splintered top meets the traveller’s eye everywhere. While
still at Masasi I wished to see this man at work, but was told that,
frightened by the rising, he had retired across the Rovuma, though
he would soon return. All subsequent inquiries as to whether the
fundi had come back met with the genuine African answer, “Bado”
(“Not yet”).
BRAZIER
Some consolation was afforded me by a brassfounder, whom I

came across in the bush near Akundonde’s. This man is the favourite
of women, and therefore no doubt of the gods; he welds the glittering
brass rods purchased at the coast into those massive, heavy rings
which, on the wrists and ankles of the local fair ones, continually give
me fresh food for admiration. Like every decent master-craftsman he
had all his tools with him, consisting of a pair of bellows, three
crucibles and a hammer—nothing more, apparently. He was quite
willing to show his skill, and in a twinkling had fixed his bellows on
the ground. They are simply two goat-skins, taken off whole, the four
legs being closed by knots, while the upper opening, intended to
admit the air, is kept stretched by two pieces of wood. At the lower
end of the skin a smaller opening is left into which a wooden tube is
stuck. The fundi has quickly borrowed a heap of wood-embers from
the nearest hut; he then fixes the free ends of the two tubes into an
earthen pipe, and clamps them to the ground by means of a bent
piece of wood. Now he fills one of his small clay crucibles, the dross
on which shows that they have been long in use, with the yellow
material, places it in the midst of the embers, which, at present are
only faintly glimmering, and begins his work. In quick alternation
the smith’s two hands move up and down with the open ends of the
bellows; as he raises his hand he holds the slit wide open, so as to let
the air enter the skin bag unhindered. In pressing it down he closes
the bag, and the air puffs through the bamboo tube and clay pipe into
the fire, which quickly burns up. The smith, however, does not keep
on with this work, but beckons to another man, who relieves him at
the bellows, while he takes some more tools out of a large skin pouch
carried on his back. I look on in wonder as, with a smooth round
stick about the thickness of a finger, he bores a few vertical holes into
the clean sand of the soil. This should not be difficult, yet the man
seems to be taking great pains over it. Then he fastens down to the
ground, with a couple of wooden clamps, a neat little trough made by
splitting a joint of bamboo in half, so that the ends are closed by the
two knots. At last the yellow metal has attained the right consistency,
and the fundi lifts the crucible from the fire by means of two sticks
split at the end to serve as tongs. A short swift turn to the left—a
tilting of the crucible—and the molten brass, hissing and giving forth
clouds of smoke, flows first into the bamboo mould and then into the
holes in the ground.
The technique of this backwoods craftsman may not be very far
advanced, but it cannot be denied that he knows how to obtain an
adequate result by the simplest means. The ladies of highest rank in
this country—that is to say, those who can afford it, wear two kinds
of these massive brass rings, one cylindrical, the other semicircular
in section. The latter are cast in the most ingenious way in the
bamboo mould, the former in the circular hole in the sand. It is quite
a simple matter for the fundi to fit these bars to the limbs of his fair
customers; with a few light strokes of his hammer he bends the
pliable brass round arm or ankle without further inconvenience to
the wearer.
SHAPING THE POT
SMOOTHING WITH MAIZE-COB
CUTTING THE EDGE

FINISHING THE BOTTOM
LAST SMOOTHING BEFORE

BURNING
FIRING THE BRUSH-PILE

LIGHTING THE FARTHER SIDE OF
THE PILE
TURNING THE RED-HOT VESSEL
NYASA WOMAN MAKING POTS AT MASASI

Pottery is an art which must always and everywhere excite the
interest of the student, just because it is so intimately connected with
the development of human culture, and because its relics are one of
the principal factors in the reconstruction of our own condition in
prehistoric times. I shall always remember with pleasure the two or
three afternoons at Masasi when Salim Matola’s mother, a slightly-
built, graceful, pleasant-looking woman, explained to me with
touching patience, by means of concrete illustrations, the ceramic art
of her people. The only implements for this primitive process were a
lump of clay in her left hand, and in the right a calabash containing
the following valuables: the fragment of a maize-cob stripped of all
its grains, a smooth, oval pebble, about the size of a pigeon’s egg, a
few chips of gourd-shell, a bamboo splinter about the length of one’s
hand, a small shell, and a bunch of some herb resembling spinach.
Nothing more. The woman scraped with the
shell a round, shallow hole in the soft, fine
sand of the soil, and, when an active young
girl had filled the calabash with water for her,
she began to knead the clay. As if by magic it
gradually assumed the shape of a rough but
already well-shaped vessel, which only wanted
a little touching up with the instruments
before mentioned. I looked out with the
MAKUA WOMAN closest attention for any indication of the use
MAKING A POT. of the potter’s wheel, in however rudimentary
SHOWS THE a form, but no—hapana (there is none). The
BEGINNINGS OF THE embryo pot stood firmly in its little
POTTER’S WHEEL
depression, and the woman walked round it in
a stooping posture, whether she was removing
small stones or similar foreign bodies with the maize-cob, smoothing
the inner or outer surface with the splinter of bamboo, or later, after
letting it dry for a day, pricking in the ornamentation with a pointed
bit of gourd-shell, or working out the bottom, or cutting the edge
with a sharp bamboo knife, or giving the last touches to the finished
vessel. This occupation of the women is infinitely toilsome, but it is
without doubt an accurate reproduction of the process in use among
our ancestors of the Neolithic and Bronze ages.
There is no doubt that the invention of pottery, an item in human
progress whose importance cannot be over-estimated, is due to
women. Rough, coarse and unfeeling, the men of the horde range
over the countryside. When the united cunning of the hunters has
succeeded in killing the game; not one of them thinks of carrying
home the spoil. A bright fire, kindled by a vigorous wielding of the
drill, is crackling beside them; the animal has been cleaned and cut
up secundum artem, and, after a slight singeing, will soon disappear
under their sharp teeth; no one all this time giving a single thought
to wife or child.
To what shifts, on the other hand, the primitive wife, and still more
the primitive mother, was put! Not even prehistoric stomachs could
endure an unvarying diet of raw food. Something or other suggested
the beneficial effect of hot water on the majority of approved but
indigestible dishes. Perhaps a neighbour had tried holding the hard
roots or tubers over the fire in a calabash filled with water—or maybe
an ostrich-egg-shell, or a hastily improvised vessel of bark. They
became much softer and more palatable than they had previously
been; but, unfortunately, the vessel could not stand the fire and got
charred on the outside. That can be remedied, thought our
ancestress, and plastered a layer of wet clay round a similar vessel.
This is an improvement; the cooking utensil remains uninjured, but
the heat of the fire has shrunk it, so that it is loose in its shell. The
next step is to detach it, so, with a firm grip and a jerk, shell and
kernel are separated, and pottery is invented. Perhaps, however, the
discovery which led to an intelligent use of the burnt-clay shell, was
made in a slightly different way. Ostrich-eggs and calabashes are not
to be found in every part of the world, but everywhere mankind has
arrived at the art of making baskets out of pliant materials, such as
bark, bast, strips of palm-leaf, supple twigs, etc. Our inventor has no
water-tight vessel provided by nature. “Never mind, let us line the
basket with clay.” This answers the purpose, but alas! the basket gets
burnt over the blazing fire, the woman watches the process of
cooking with increasing uneasiness, fearing a leak, but no leak
appears. The food, done to a turn, is eaten with peculiar relish; and
the cooking-vessel is examined, half in curiosity, half in satisfaction
at the result. The plastic clay is now hard as stone, and at the same
time looks exceedingly well, for the neat plaiting of the burnt basket
is traced all over it in a pretty pattern. Thus, simultaneously with
pottery, its ornamentation was invented.
Primitive woman has another claim to respect. It was the man,
roving abroad, who invented the art of producing fire at will, but the
woman, unable to imitate him in this, has been a Vestal from the
earliest times. Nothing gives so much trouble as the keeping alight of
the smouldering brand, and, above all, when all the men are absent
from the camp. Heavy rain-clouds gather, already the first large
drops are falling, the first gusts of the storm rage over the plain. The
little flame, a greater anxiety to the woman than her own children,
flickers unsteadily in the blast. What is to be done? A sudden thought
occurs to her, and in an instant she has constructed a primitive hut
out of strips of bark, to protect the flame against rain and wind.
This, or something very like it, was the way in which the principle
of the house was discovered; and even the most hardened misogynist
cannot fairly refuse a woman the credit of it. The protection of the
hearth-fire from the weather is the germ from which the human
dwelling was evolved. Men had little, if any share, in this forward
step, and that only at a late stage. Even at the present day, the
plastering of the housewall with clay and the manufacture of pottery
are exclusively the women’s business. These are two very significant
survivals. Our European kitchen-garden, too, is originally a woman’s
invention, and the hoe, the primitive instrument of agriculture, is,
characteristically enough, still used in this department. But the
noblest achievement which we owe to the other sex is unquestionably
the art of cookery. Roasting alone—the oldest process—is one for
which men took the hint (a very obvious one) from nature. It must
have been suggested by the scorched carcase of some animal
overtaken by the destructive forest-fires. But boiling—the process of
improving organic substances by the help of water heated to boiling-
point—is a much later discovery. It is so recent that it has not even
yet penetrated to all parts of the world. The Polynesians understand
how to steam food, that is, to cook it, neatly wrapped in leaves, in a
hole in the earth between hot stones, the air being excluded, and
(sometimes) a few drops of water sprinkled on the stones; but they
do not understand boiling.
To come back from this digression, we find that the slender Nyasa
woman has, after once more carefully examining the finished pot,
put it aside in the shade to dry. On the following day she sends me
word by her son, Salim Matola, who is always on hand, that she is
going to do the burning, and, on coming out of my house, I find her
already hard at work. She has spread on the ground a layer of very
dry sticks, about as thick as one’s thumb, has laid the pot (now of a
yellowish-grey colour) on them, and is piling brushwood round it.
My faithful Pesa mbili, the mnyampara, who has been standing by,
most obligingly, with a lighted stick, now hands it to her. Both of
them, blowing steadily, light the pile on the lee side, and, when the
flame begins to catch, on the weather side also. Soon the whole is in a
blaze, but the dry fuel is quickly consumed and the fire dies down, so
that we see the red-hot vessel rising from the ashes. The woman
turns it continually with a long stick, sometimes one way and
sometimes another, so that it may be evenly heated all over. In
twenty minutes she rolls it out of the ash-heap, takes up the bundle
of spinach, which has been lying for two days in a jar of water, and
sprinkles the red-hot clay with it. The places where the drops fall are
marked by black spots on the uniform reddish-brown surface. With a
sigh of relief, and with visible satisfaction, the woman rises to an
erect position; she is standing just in a line between me and the fire,
from which a cloud of smoke is just rising: I press the ball of my
camera, the shutter clicks—the apotheosis is achieved! Like a
priestess, representative of her inventive sex, the graceful woman
stands: at her feet the hearth-fire she has given us beside her the
invention she has devised for us, in the background the home she has
built for us.
At Newala, also, I have had the manufacture of pottery carried on
in my presence. Technically the process is better than that already
described, for here we find the beginnings of the potter’s wheel,
which does not seem to exist in the plains; at least I have seen
nothing of the sort. The artist, a frightfully stupid Makua woman, did
not make a depression in the ground to receive the pot she was about
to shape, but used instead a large potsherd. Otherwise, she went to
work in much the same way as Salim’s mother, except that she saved
herself the trouble of walking round and round her work by squatting
at her ease and letting the pot and potsherd rotate round her; this is
surely the first step towards a machine. But it does not follow that
the pot was improved by the process. It is true that it was beautifully
rounded and presented a very creditable appearance when finished,
but the numerous large and small vessels which I have seen, and, in
part, collected, in the “less advanced” districts, are no less so. We
moderns imagine that instruments of precision are necessary to
produce excellent results. Go to the prehistoric collections of our
museums and look at the pots, urns and bowls of our ancestors in the
dim ages of the past, and you will at once perceive your error.
MAKING LONGITUDINAL CUT IN
BARK
DRAWING THE BARK OFF THE LOG
REMOVING THE OUTER BARK

BEATING THE BARK
WORKING THE BARK-CLOTH AFTER BEATING, TO MAKE IT

SOFT
MANUFACTURE OF BARK-CLOTH AT NEWALA

To-day, nearly the whole population of German East Africa is
clothed in imported calico. This was not always the case; even now in
some parts of the north dressed skins are still the prevailing wear,
and in the north-western districts—east and north of Lake
Tanganyika—lies a zone where bark-cloth has not yet been
superseded. Probably not many generations have passed since such
bark fabrics and kilts of skins were the only clothing even in the
south. Even to-day, large quantities of this bright-red or drab
material are still to be found; but if we wish to see it, we must look in
the granaries and on the drying stages inside the native huts, where
it serves less ambitious uses as wrappings for those seeds and fruits
which require to be packed with special care. The salt produced at
Masasi, too, is packed for transport to a distance in large sheets of
bark-cloth. Wherever I found it in any degree possible, I studied the
process of making this cloth. The native requisitioned for the
purpose arrived, carrying a log between two and three yards long and
as thick as his thigh, and nothing else except a curiously-shaped
mallet and the usual long, sharp and pointed knife which all men and
boys wear in a belt at their backs without a sheath—horribile dictu!
[51]
Silently he squats down before me, and with two rapid cuts has
drawn a couple of circles round the log some two yards apart, and
slits the bark lengthwise between them with the point of his knife.
With evident care, he then scrapes off the outer rind all round the
log, so that in a quarter of an hour the inner red layer of the bark
shows up brightly-coloured between the two untouched ends. With
some trouble and much caution, he now loosens the bark at one end,
and opens the cylinder. He then stands up, takes hold of the free
edge with both hands, and turning it inside out, slowly but steadily
pulls it off in one piece. Now comes the troublesome work of
scraping all superfluous particles of outer bark from the outside of
the long, narrow piece of material, while the inner side is carefully
scrutinised for defective spots. At last it is ready for beating. Having
signalled to a friend, who immediately places a bowl of water beside
him, the artificer damps his sheet of bark all over, seizes his mallet,
lays one end of the stuff on the smoothest spot of the log, and
hammers away slowly but continuously. “Very simple!” I think to
myself. “Why, I could do that, too!”—but I am forced to change my
opinions a little later on; for the beating is quite an art, if the fabric is
not to be beaten to pieces. To prevent the breaking of the fibres, the
stuff is several times folded across, so as to interpose several
thicknesses between the mallet and the block. At last the required
state is reached, and the fundi seizes the sheet, still folded, by both
ends, and wrings it out, or calls an assistant to take one end while he
holds the other. The cloth produced in this way is not nearly so fine
and uniform in texture as the famous Uganda bark-cloth, but it is
quite soft, and, above all, cheap.
Now, too, I examine the mallet. My craftsman has been using the
simpler but better form of this implement, a conical block of some
hard wood, its base—the striking surface—being scored across and
across with more or less deeply-cut grooves, and the handle stuck
into a hole in the middle. The other and earlier form of mallet is
shaped in the same way, but the head is fastened by an ingenious
network of bark strips into the split bamboo serving as a handle. The
observation so often made, that ancient customs persist longest in
connection with religious ceremonies and in the life of children, here
finds confirmation. As we shall soon see, bark-cloth is still worn
during the unyago,[52] having been prepared with special solemn
ceremonies; and many a mother, if she has no other garment handy,
will still put her little one into a kilt of bark-cloth, which, after all,
looks better, besides being more in keeping with its African
surroundings, than the ridiculous bit of print from Ulaya.
MAKUA WOMEN

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eTextbook 978-0077837280 Ecology:

Concepts and Applications 7th Edition

8.2 Mate Choice and Resource Provisioning

8.4 Sociality 184

Concepts 258 Applications: Competition between Native

Section IV Models 317

15.3 Evolution of Mutualism 344 17.2 Indirect Interactions 376

Chapter 20 Succession and Stability 435

Concepts 435 Structure 472

Manipulating Island Area 493 El Niño and Marine Populations 511

ecology and includes chapters on landscape, geographic,

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

historical events related to the subject; others pre- moL37282_ch05_

sent an example of an ecological process. All attempt 99

students with key information where they Birds leave the

of on the y-axis creates a tio

pi- Dall sheep surviving their

100 wolves and die

Confirming Pag Confirming Pag

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

use Connect’s robust reporting features to generate powerful

Save Time with Auto-Graded

McGraw-Hill LearnSmart® is the only adaptive learning

What You’ve Only Imagined

Annual Editions: Environment 2015 Acknowledgments

Jamal Bittar The University of Toledo Jerry Baskin University of Kentucky

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

1.1 Discuss the concept of environment as it pertains to

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

1.1 Overview of Ecology

Ecologists study environmental relationships ranging Interactions

distribution and abundance of species, population growth and

Concept 1.1 Review

Cape May warbler Blackburnian warbler Black-throated

Old needles and bare Bare or lichen-covered lower

American redstarts breed across

American redstarts winter

Figure 1.4 A male American redstart, Setophaga ruticilla.

Nearly half a century after Robert MacArthur studied the

Forest Canopy Research:

Gondola housing scientists can Crane operator can swing crane

Lake profile Chestnut decline Present Pollen

Concept 1.2 Review

1. How were the warbler studies of Robert MacArthur and

Investigating the Evidence 1

The Scientific Method—Questions and Hypotheses

LEARNING OUTCOMES Information: Testing a hypothesis,

Newala, too, suffers from the distance of its water-supply—at least

The water-supply of New Newala is in the bottom of the valley,

NATIVE PATH THROUGH THE MAKONDE BUSH, NEAR

MAKONDE LOCK AND KEY AT JUMBE CHAURO

MODE OF INSERTING THE KEY

With no small pride first one householder and then a second

Some consolation was afforded me by a brassfounder, whom I

SMOOTHING WITH MAIZE-COB

CUTTING THE EDGE

LAST SMOOTHING BEFORE

FIRING THE BRUSH-PILE

TURNING THE RED-HOT VESSEL

NYASA WOMAN MAKING POTS AT MASASI

DRAWING THE BARK OFF THE LOG

REMOVING THE OUTER BARK

WORKING THE BARK-CLOTH AFTER BEATING, TO MAKE IT