Ecology: Concepts and Applications

593 Pages·2015·132.143 MB·English

by  Manuel Molles

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Ecology
SEVENTH EDITION
Concepts and Applications
Manuel C. Molles Jr.
University of New Mexico
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ECOLOGY: CONCEPTS AND APPLICATIONS, SEVENTH EDITION
Published by McGraw-Hill Education, 2 Penn Plaza, New York, NY 10121. Copyright © 2016
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Library of Congress Cataloging-in-Publication Data
Molles, Manuel C., Jr., 1948-
Ecology : concepts and applications / Manuel C. Molles, Jr., University of New Mexico.
—Seventh edition.
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About the Author
Manuel C. Molles Jr.
is an emeritus Professor of Biology at the
U niversity of New Mexico, where he has been a member of the faculty and curator in
the Museum of Southwestern Biology since 1975 and where he continues to write and
conduct ecological research. He received his B.S. from Humboldt State University and
his Ph.D. from the Department of Ecology and Evolutionary Biology at the University
of Arizona. Seeking to broaden his geographic perspective, he has taught and conducted
ecological research in Latin America, the Caribbean, and Europe. He was awarded a
Fulbright Research Fellowship to conduct research on river ecology in Portugal and has
held visiting professor appointments in the Department of Zoology at the University
of Coimbra, Portugal, in the Laboratory of Hydrology at the Polytechnic University of
Madrid, Spain, and at the University of Montana’s Flathead Lake Biological Station.
Originally trained as a marine ecologist and fisheries biologist, the author has
worked mainly on river and riparian ecology at the University of New Mexico. His
research has covered a wide range of ecological levels, including behavioral ecology,
population biology, community ecology, ecosystem ecology, biogeography of stream
insects, and the influence of a large-scale climate system (El Niño) on the dynamics
of southwestern river and riparian ecosystems. His current research concerns the influ-
ence of climate change and climatic variability on the dynamics of populations and
communities along steep gradients of temperature and moisture in the mountains of
the Southwest. Throughout his career, Dr. Molles has attempted to combine research,
teaching, and service, involving undergraduate as well as graduate students in his ongo-
ing projects. At the University of New Mexico, he has taught a broad range of lower
division, upper division, and graduate courses, including Principles of Biology, Evolu-
tion and Ecology, Stream Ecology, Limnology and Oceanography, Marine Biology, and
Community and Ecosystem Ecology. He has taught courses in Global Change and River
Ecology at the University of Coimbra, Portugal, and General Ecology and Groundwater
and Riparian Ecology at the Flathead Lake Biological Station. Dr. Manuel Molles was
named Teacher of the Year by the University of New Mexico for 1995–1996 and Potter
Chair in Plant Ecology in 2000. In 2014, he received the Eugene P. Odum Award from
the Ecological Society of America based on his “ability to relate basic ecological prin-
ciples to human affairs through teaching, outreach and mentoring activities.”
Dedication
To Mary Anne
and
Keena
iii
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Brief Contents
1 Introduction to Ecology: Historical Foundations and Developing Frontiers 1
Section N atural History and Evolution 11
I
2 Life on Land 11
3 Life in Water 45
4 Population Genetics and Natural Selection 77
Section A daptations to the Environment 99
II
5 Temperature Relations 99
6 Water Relations 125
7 Energy and Nutrient Relations 149
8 Social Relations 173
Section Population Ecology 198
III
9 Population Distribution and Abundance 198
10 Population Dynamics 218
11 Population Growth 241
12 Life Histories 258
Section I nteractions 282
IV
13 Competition 282
14 Exploitative Interactions: Predation, Herbivory, Parasitism, and Disease 303
15 Mutualism 331
Section C ommunities and Ecosystems 352
V
16 Species Abundance and Diversity 352
17 S pecies Interactions and Community Structure 372
18 P rimary and Secondary Production 392
19 N utrient Cycling and Retention 414
20 Succession and Stability 435
Section Large-Scale Ecology 460
VI
21 L andscape Ecology 460
22 G eographic Ecology 484
23 G lobal Ecology 506
Appendix Statistical Tables 529
iv
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Contents
3
Preface xiii Chapter Life in Water 45
Concepts 45
1
Chapter Introduction to Ecology: Historical 3.1 The Hydrologic Cycle 46
Foundations and Developing Concept 3.1 Review 46
Frontiers 1 3.2 The Natural History of Aquatic Environments 46
The Oceans 47
Concepts 1
Life in Shallow Marine Waters: Kelp Forests
1.1 Overview of Ecology 2
and Coral Gardens 51
Concept 1.1 Review 3 Investigating the Evidence 3: Determining the Sample
1.2 Sampling Ecological Research 3 Median 52
The Ecology of Forest Birds: Old Tools and New 4 Marine Shores: Life Between High and Low Tides 55
Forest Canopy Research: A Physical and Scientific Frontier 6 Transitional Environments: Estuaries, Salt Marshes,
Climatic and Ecological Change: Past and Future 7 Mangrove Forests, and Freshwater Wetlands 58
Concept 1.2 Review 8 Rivers and Streams: Life Blood and Pulse
of the Land 63
Investigating the Evidence 1: The Scientific Method—
Questions and Hypotheses 9 Lakes: Small Seas 67
Concept 3.2 Review 72
I Applications: Biological Integrity—Assessing the Health
Section
of Aquatic Systems 72
NATURAL HISTORY AND EVOLUTION Number of Species and Species Composition 73
Trophic Composition 73
2 Fish Abundance and Condition 73
Chapter Life on Land 11
A Test 73
Concepts 11
Terrestrial Biomes 12
4
2.1 Large-Scale Patterns of Climatic Variation 13 Chapter Population Genetics and Natural
Temperature, Atmospheric Circulation, and Precipitation 13 Selection 77
Climate Diagrams 15
Concepts 77
Concept 2.1 Review 16
4.1 Variation Within Populations 79
2.2 Soil: The Foundation of Terrestrial Biomes 16
Variation in a Widely Distributed Plant 80
Investigating the Evidence 2: Determining the Sample
Variation in Alpine Fish Populations 80
Mean 18
Concept 4.1 Review 82
Concept 2.2 Review 19
4.2 Hardy-Weinberg Principle 83
2.3 Natural History and Geography of Biomes 19
Calculating Gene Frequencies 83
Tropical Rain Forest 20
Concept 4.2 Review 85
Tropical Dry Forest 21
Tropical Savanna 23 4.3 The Process of Natural Selection 85
Desert 25 Stabilizing Selection 85
Mediterranean Woodland and Shrubland 27 Directional Selection 86
Temperate Grassland 30 Disruptive Selection 86
Temperate Forest 31 Concept 4.3 Review 87
Boreal Forest 34 4.4 Evolution by Natural Selection 87
Tundra 35 Heritability: Essential for Evolution 87
Mountains: Islands in the Sky 38 Investigating the Evidence 4: Variation in Data 88
Concept 2.3 Review 41 Directional Selection: Adaptation by Soapberry Bugs
Applications: Climatic Variation and the Palmer Drought to New Host Plants 89
Severity Index 41 Concept 4.4 Review 92
v
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vi Contents
4.5 Change Due to Chance 92 6.2 Water Regulation on Land 131
Evidence of Genetic Drift in Chihuahua Spruce 92 Water Acquisition by Animals 131
Genetic Variation in Island Populations 93 Water Acquisition by Plants 133
Genetic Diversity and Butterfly Extinctions 94 Water Conservation by Plants and Animals 134
Concept 4.5 Review 95 Investigating the Evidence 6: Sample Size 136
Applications: Evolution and Agriculture 95 Dissimilar Organisms with Similar Approaches
to Desert Life 138
Evolution of Herbicide Resistance in Weeds 96
Two Arthropods with Opposite Approaches
II to Desert Life 140
Section Concept 6.2 Review 142
6.3 Water and Salt Balance in Aquatic
ADAPTATIONS TO THE ENVIRONMENT
Environments 142
5 Marine Fish and Invertebrates 142
Chapter Temperature Relations 99
Freshwater Fish and Invertebrates 143
Concepts 99 Concept 6.3 Review 144
5.1 Microclimates 100 Applications: Using Stable Isotopes to Study Water Uptake
by Plants 144
Altitude 100
Aspect 101 Stable Isotope Analysis 145
Vegetation 101 Using Stable Isotopes to Identify Plant Water
Color of the Ground 101 Sources 146
Presence of Boulders and Burrows 102
7
Aquatic Temperatures 102 Chapter Energy and Nutrient Relations 149
Concept 5.1 Review 103
Concepts 149
5.2 Evolutionary Trade-Offs 103
7.1 Photosynthetic Autotrophs 151
The Principle of Allocation 104
Concept 5.2 Review 104 The Solar-Powered Biosphere 151
Concept 7.1 Review 155
5.3 Temperature and Performance of Organisms 105
7.2 Chemosynthetic Autotrophs 155
Investigating the Evidence 5: Laboratory Experiments 106
Extreme Temperatures and Photosynthesis 107 Concept 7.2 Review 155
Temperature and Microbial Activity 108 7.3 Heterotrophs 155
Concept 5.3 Review 109 Chemical Composition and Nutrient Requirements 156
5.4 Regulating Body Temperature 109 Concept 7.3 Review 163
Balancing Heat Gain against Heat Loss 109 7.4 Energy Limitation 163
Temperature Regulation by Plants 110 Photon Flux and Photosynthetic Response Curves 163
Temperature Regulation by Ectothermic Animals 112 Food Density and Animal Functional Response 164
Temperature Regulation by Endothermic Animals 114 Concept 7.4 Review 165
Temperature Regulation by Thermogenic Plants 118 7.5 Optimal Foraging Theory 165
Concept 5.4 Review 119
Testing Optimal Foraging Theory 166
5.5 Surviving Extreme Temperatures 119 Optimal Foraging by Plants 167
Inactivity 119 Investigating the Evidence 7: Scatter Plots and the
Reducing Metabolic Rate 120 Relationship between Variables 168
Hibernation by a Tropical Species 120 Concept 7.5 Review 169
Concept 5.5 Review 121 Applications: Bioremediation—Using the Trophic
Applications: Local Extinction of a Land Snail in an Urban Diversity of Bacteria to Solve Environmental
Heat Island 122 Problems 169
Leaking Underground Storage Tanks 169
6
Chapter Water Relations 125 Cyanide and Nitrates in Mine Spoils 170
Concepts 125 8
Chapter Social Relations 173
6.1 Water Availability 127
Concepts 173
Water Content of Air 127
Water Movement in Aquatic Environments 128 8.1 Mate Choice versus Predation 175
Water Movement between Soils and Plants 129 Mate Choice and Sexual Selection in Guppies 176
Concept 6.1 Review 130 Concept 8.1 Review 179
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Contents vii
8.2 M ate Choice and Resource Provisioning 179 Chapter 10 Population Dynamics 218
Concept 8.2 Review 182
Concepts 218
8.3 N onrandom Mating in a Plant Population 182
Concept 8.3 Review 184 10.1 Dispersal 220
Dispersal of Expanding Populations 220
8.4 Sociality 184
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
Dispersal in Rivers and Streams 223
Regression Analysis 188
Concept 10.1 Review 224
Concept 8.4 Review 191
10.2 Metapopulations 224
8.5 Eusociality 191
A Metapopulation of an Alpine Butterfly 225
Eusocial Species 191
Dispersal Within a Metapopulation of Lesser Kestrels 226
Evolution of Eusociality 193
Concept 10.2 Review 227
Concept 8.5 Review 195
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
Constant Rates of Survival 229
of Behavior 195
High Mortality Among the Young 230
Three Types of Survivorship Curves 230
Concept 10.3 Review 231
III
Section 10.4 Age Distribution 231
Contrasting Tree Populations 231
POPULATION ECOLOGY
A Dynamic Population in a Variable Climate 232
Concept 10.4 Review 233
9
Chapter 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
Concept 10.5 Review 237
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 11
Chapter Population Growth 241
Gradient 203
Concept 9.1 Review 204 Concepts 241
9.2 Patterns on Small Scales 204 11.1 G eometric 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 O rganism 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
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viii Contents
12 Investigating the Evidence 13: Field Experiments 299
Chapter Life Histories 258
Concept 13.4 Review 300
Concepts 258
Applications: Competition between Native
12.1 O ffspring Number Versus Size 259 and Invasive Species 300
Egg Size and Number in Fish 260
Seed Size and Number in Plants 262 14
Chapter Exploitative Interactions: Predation,
Seed Size and Seedling Performance 263
Herbivory, Parasitism, and
Concept 12.1 Review 265
Disease 303
12.2 A dult Survival and Reproductive Allocation 266
Life History Variation Among Species 266 Concepts 303
Life History Variation Within Species 267
14.1 Complex Interactions 304
Concept 12.2 Review 270
Parasites and Pathogens that Manipulate Host
12.3 Life History Classification 270 Behavior 304
r and K Selection 270 The Entangling of Exploitation with Competition 307
Plant Life Histories 271 Concept 14.1 Review 308
Investigating the Evidence 12: A Statistical Test 14.2 Exploitation and Abundance 308
for Distribution Pattern 272
A Herbivorous Stream Insect and Its Algal Food 308
Opportunistic, Equilibrium, and Periodic Life
Bats, Birds, and Herbivory in a Tropical Forest 309
Histories 274
A Pathogenic Parasite, a Predator, and Its Prey 311
Lifetime Reproductive Effort and Relative Offspring Size:
Concept 14.2 Review 312
Two Central Variables? 275
14.3 Dynamics 312
Concept 12.3 Review 276
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
IV Models 317
Section
Concept 14.3 Review 319
INTERACTIONS 14.4 Refuges 320
Refuges and Host Persistence in Laboratory
Chapter 13 Competition 282 and Mathematical Models 320
Exploited Organisms and Their Wide Variety
Concepts 282 of “Refuges” 321
13.1 Intraspecific Competition 284 Concept 14.4 Review 323
Intraspecific Competition Among Plants 284 14.5 Ratio-Dependent Models of Functional Response 323
Intraspecific Competition Among Planthoppers 285 Alternative Model for Trophic Ecology 324
Interference Competition Among Terrestrial Isopods 285 Evidence for Ratio-Dependent Predation 324
Concept 13.1 Review 286 Concept 14.5 Review 326
13.2 Competitive Exclusion and Niches 286 Applications: The Value of Pest Control by Bats:
The Feeding Niches of Darwin’s Finches 286 A Case Study 327
The Habitat Niche of a Salt Marsh Grass 288
Concept 13.2 Review 289 15
Chapter Mutualism 331
13.3 Mathematical and Laboratory Models 289
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
A Temperate Plant Protection Mutualism 340
Niches and Competition Among Plants 293
Concept 15.1 Review 341
Niche Overlap and Competition between Barnacles 293
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
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Contents ix
15.3 Evolution of Mutualism 344 17.2 Indirect Interactions 376
Investigating the Evidence 15: Confidence Intervals 345 Indirect Commensalism 376
Facultative Ant-Plant Protection Mutualisms 347 Apparent Competition 376
Concept 15.3 Review 348 Concept 17.2 Review 378
Applications: Mutualism and Humans 348 17.3 Keystone Species 378
Guiding Behavior 348 Food Web Structure and Species Diversity 379
Experimental Removal of Sea Stars 380
V Snail Effects on Algal Diversity 381

Section Fish as Keystone Species in River Food Webs 383
Investigating the Evidence 17: Using Confidence Intervals
COMMUNITIES AND ECOSYSTEMS
to Compare Populations 384
16 Concept 17.3 Review 386
Chapter Species Abundance
17.4 M utualistic Keystones 386
and Diversity 352
A Cleaner Fish as a Keystone Species 386
Concepts 352 Seed Dispersal Mutualists as Keystone Species 387
Concept 17.4 Review 388
16.1 Species Abundance 354
The Lognormal Distribution 354 Applications: Human Modification of Food Webs 388
Concept 16.1 Review 355 The Empty Forest: Hunters and Tropical Rain Forest
Animal Communities 388
16.2 Species Diversity 355
Ants and Agriculture: Keystone Predators for Pest
A Quantitative Index of Species Diversity 355
Control 389
Rank-Abundance Curves 356
Concept 16.2 Review 357
18
16.3 Environmental Complexity 357 Chapter Primary and Secondary
Forest Complexity and Bird Species Diversity 358 Production 392
Investigating the Evidence 16: Estimating the Number
of Species in Communities 359 Concepts 392
Niches, Heterogeneity, and the Diversity of Algae and 18.1 Patterns of Terrestrial Primary Production 394
Plants 360 Actual Evapotranspiration and Terrestrial Primary
The Niches of Algae and Terrestrial Plants 360 Production 394
Complexity in Plant Environments 361 Soil Fertility and Terrestrial Primary Production 395
Soil and Topographic Heterogeneity and the Diversity Concept 18.1 Review 396
of Tropical Forest Trees 361
18.2 Patterns of Aquatic Primary Production 396
Algal and Plant Species Diversity and Increased Nutrient
Patterns and Models 396
Availability 363
Whole Lake Experiments on Primary
Nitrogen Enrichment and Ectomycorrhizal Fungus
Production 397
Diversity 363
Global Patterns of Marine Primary Production 397
Concept 16.3 Review 364
Concept 18.2 Review 398
16.4 Disturbance and Diversity 364
18.3 Primary Producer Diversity 399
The Nature and Sources of Disturbance 364
Terrestrial Plant Diversity and Primary Production 399
The Intermediate Disturbance Hypothesis 364
Algal Diversity and Aquatic Primary Production 400
Disturbance and Diversity in the Intertidal Zone 365
Concept 18.3 Review 400
Disturbance and Diversity in Temperate Grasslands 365
18.4 Consumer Influences 401
Concept 16.4 Review 367
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
17
Chapter Species Interactions Concept 18.4 Review 405
and C ommunity Structure 372 18.5 Secondary Production 405
Investigating the Evidence 18: Comparing Two Populations
Concepts 372
with the t-Test 406
17.1 Community Webs 374 A Trophic Dynamic View of Ecosystems 406
Detailed Food Webs Reveal Great Complexity 374 Linking Primary Production
Strong Interactions and Food Web Structure 374 and Secondary Production 408
Concept 17.1 Review 375 Concept 18.5 Review 409
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