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ECOLOGY

Published by McGraw Hill LLC, 1325 Avenue of the Americas, New York, NY 10019. Copyright
© 2022 by McGraw Hill LLC. All rights reserved. Printed in the United States of America. No part
of this publication may be reproduced or distributed in any form or by any means, or stored in a
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ISBN 978-1-265-28633-0
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Cover Image: Anna A. Sher

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 About the Authors
Anna A. Sher is a full professor in the Department of Biological Sciences at the Uni-
versity of Denver, where she has been faculty since 2003. Until 2010 she held this position jointly
with the Denver Botanic Gardens as the Director of Research and Conservation. As a student, she
was a double major in Biology and Art at Earlham College, where she has also taught ecology, and
was the co-leader of the Earlham Study Abroad Kenya Program. She received her PhD from the
University of New Mexico, where she also taught botany as a visiting lecturer. As a postdoctoral
researcher, Dr. Sher was awarded a Fulbright postdoctoral research fellowship to conduct research
on plant interactions in Israel at Ben Gurion University’s Mitrani Department of Desert Ecology,
and she also studied the ecology of an invasive grass at the University of California, Davis. She has
also been a visiting professor at the University of Otago, Dunedin, New Zealand.
Dr. Sher’s primary research focus has been on the ecological dynamics associated with
the removal of invasive riparian plants. She is known as a leading expert in the ecology of
Tamarix, a dominant exotic tree, and she was the lead editor of the first book exclusively on
the topic. Her research interests and publications have spanned several areas within ecol-
ogy, including not only restoration ecology, competition, and invasive species ecology, but
also interactions between plants and soil chemistry, mycorrhizae, insect diversity and trophic
cascades, ethnobotany, phenology, climate change, and rare species conservation. She is also
lead author of the textbook series An Introduction to Conservation Biology (Oxford University Courtesy of Anna Sher
Press). Dr. Sher has a particular interest in quantitative ecological methods, with her lab
specializing in multivariate methods and spatial models at both individual organism and regional scales. She is currently principal
investigator of a National Science Foundation award to investigate the human dimension of the restoration of damaged ecosys-
tems, and she has been a TEDx speaker on the way ecosystems can teach us how to solve human problems.
Above all, Dr. Sher loves to teach and mentor students doing research at both undergraduate and graduate levels.

Manuel C. Molles Jr. is an emeritus Professor of Biology at the ­University of New Mexico, where he has been a
member of the faculty and curator in the Museum of Southwestern ­Biology since 1975. He received his BS from Humboldt State
­University and his PhD 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 con-
duct 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 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 interests focus on the influence of climate change and cli-
matic 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 ongoing projects.
At the University of New Mexico, he taught a broad range of lower division, upper division, and graduate
courses, including Principles of Biology, Evolution and Ecology, Stream Ecology, Limnology and Oceanog-
raphy, 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 principles to human affairs through teaching, outreach and mentoring activities.”
Courtesy of Manuel Molles
Design elements: Anna A. Sher
iii
 Dedication
To the Sher Lab and the whole next generation
of ecologists, who inspire me to do this work.
Also, I dedicate this edition to my co-author
and mentor, Manuel.
—Anna A. Sher
 Brief Contents
1 Introduction to Ecology: Historical Foundations and Developing Frontiers 1

Section Natural History and Evolution 11

I 2 Life on Land 11
3 Life in Water 44
4 Population Genetics and Natural Selection 78

Section Adaptations to the Environment 101

II 5
6
7
Temperature Relations 101
Water Relations 127
Energy and Nutrient Relations 149
8 Social Relations 172

Section Population Ecology 196

III 9
10
11
Population Distribution and Abundance
Population Dynamics 215
Population Growth 237
196

12 Life Histories 254

Section Interactions 277

IV 13 Species Interactions and Competition 277

14 Exploitative Interactions: Predation, Herbivory, Parasitism, and Disease 299
15 Mutualism 325

Section Communities and Ecosystems 345

V 16
17
18
Species Abundance and Diversity 345
Species Interactions and Community Structure
Primary and Secondary Production 383
365

19 Nutrient Cycling and Retention 403

20 Succession and Stability 423

Section Large-Scale Ecology 445

VI 21 Landscape Ecology 445

22 Geographic Ecology 468
23 Global Ecology 490

Appendix A   Investigating the Evidence 514

Appendix B   Statistical Tables 541
Appendix C   Abbreviations Used in This Text 545
Appendix D   Global Biomes 547

Design elements: Anna A. Sher

v
Contents
Preface xiii
Chapter 3 Life in Water 44
Chapter 1 Introduction to Ecology: Concepts 44

Historical Foundations and Aquatic Biomes and How They Differ 45

Developing Frontiers 1 3.1 Water Cycling 47
The Hydrologic Cycle 47
Concepts 1
The Effects of Wind and Temperature 47
1.1 Overview of Ecology 2 Concept 3.1 Review 48
Concept 1.1 Review 3 3.2 The Natural History of Aquatic Environments 49
1.2 Sampling Ecological Research 3 The Oceans 49
Climatic and Ecological Change: Past and Future 7 Life in Shallow Marine Waters: Kelp Forests and Coral
Concept 1.2 Review 9 Gardens 54
Applications: Ecology Can Inform Environmental Law Marine Shores: Life Between High and Low Tides 57
and Policy 9 Transitional Environments: Estuaries, Salt Marshes,
­Mangrove Forests, and Freshwater Wetlands 59
Rivers and Streams: Life Blood and Pulse of the Land 64

Section I Lakes: Small Seas 69

Concept 3.2 Review 73
NATURAL HISTORY AND EVOLUTION Applications: Biological Integrity—Assessing the Health of
Aquatic Systems 73
Chapter 2 Life on Land 11 Number of Species and Species Composition 74
Trophic Composition 74
Concepts 11
Fish Abundance and Condition 74
Terrestrial Biomes and the Importance A Test 74
of Plants 12
2.1 Large-Scale Patterns of Climatic Variation 14
Temperature, Atmospheric Circulation, and Chapter 4 Population Genetics and Natural
Precipitation 14 Selection 78
Climate Diagrams 16
Concepts 78
Concept 2.1 Review 16
4.1 Variation Within Populations 81
2.2 Other Factors That Shape Terrestrial
Biomes 17 Variation in a Widely Distributed Plant 81
Concept 2.2 Review 19 Variation in Alpine Fish Populations 83
Concept 4.1 Review 84
2.3 Natural History and Geography of Biomes 19
4.2 Hardy-Weinberg Principle 84
Tropical Rain Forest 19
Tropical Dry Forest 21 Calculating Gene Frequencies 84
Tropical Savanna 22 Concept 4.2 Review 86
Desert 25 4.3 The Process of Natural Selection 87
Woodland and Shrubland 27 Stabilizing Selection 87
Temperate Grassland 29 Directional Selection 87
Temperate Forest 31 Disruptive Selection 87
Boreal Forest 32 Concept 4.3 Review 88
Tundra 35 4.4 Evolution by Natural Selection 88
Mountains: A Diversity of Biomes 37 Heritability: Essential for Evolution 89
Concept 2.3 Review 40 Directional Selection: Adaptation by Soapberry Bugs to
Applications: Finer Scale Climatic Variation over Time New Host Plants 90
and Space 40 Concept 4.4 Review 92

vi
 Contents vii

4.5 Change due to Chance 92 Water Acquisition by Plants 135

Evidence of Genetic Drift in Island Crickets 93 Water Conservation by Plants and Animals 136
Genetic Diversity and Butterfly Extinctions 95 Dissimilar Organisms with Similar Approaches to Desert
Concept 4.5 Review 96 Life 139
Applications: Evolution and Agriculture 96 Two Arthropods with Opposite Approaches to Desert
Life 140
Evolution of Herbicide Resistance in Weeds 97
Concept 6.2 Review 141
6.3 Water and Salt Balance in Aquatic Environments 143

Section II Marine Fish and Invertebrates 143

Freshwater Fish and Invertebrates 144
ADAPTATIONS TO THE ENVIRONMENT Concept 6.3 Review 144
Applications: Using Stable Isotopes to Study Water Uptake by
Chapter 5 Temperature Relations 101 Plants 145
Stable Isotope Analysis 146
Concepts 101
Using Stable Isotopes to Identify Plant Water Sources 146
5.1 Microclimates 102
Altitude 103
Aspect 103
Chapter 7 Energy and Nutrient Relations 149
Vegetation 103 Concepts 149
Color of the Ground 103 7.1 Photosynthetic Autotrophs 150
Presence of Boulders and Burrows 104
The Solar-Powered Biosphere 150
Aquatic Temperatures 104
Concept 7.1 Review 153
Concept 5.1 Review 105
7.2 Chemosynthetic Autotrophs 154
5.2 Evolutionary Trade-Offs 105
Concept 7.2 Review 156
The Principle of Allocation 105
7.3 Heterotrophs 156
Concept 5.2 Review 106
Chemical Composition and Nutrient Requirements 156
5.3 Temperature and Performance of Organisms 106
Concept 7.3 Review 162
Extreme Temperatures and Photosynthesis 108
7.4 Energy Limitation 163
Temperature and Microbial Activity 109
Concept 5.3 Review 110 Photon Flux and Photosynthetic Response Curves 163
Food Density and Animal Functional Response 164
5.4 Regulating Body Temperature 110
Concept 7.4 Review 165
Balancing Heat Gain Against Heat Loss 110
7.5 Optimal Foraging Theory 165
Temperature Regulation by Plants 111
Temperature Regulation by Ectothermic Animals 113 Testing Optimal Foraging Theory 166
Temperature Regulation by Endothermic Animals 115 Optimal Foraging by Plants 167
Temperature Regulation by Thermogenic Plants 119 Concept 7.5 Review 168
Concept 5.4 Review 120 Applications: Bioremediation—Using the Trophic
­Diversity of Bacteria to Solve Environmental
5.5 Surviving Extreme Temperatures 120
Problems 168
Inactivity 120
Leaking Underground Storage Tanks 169
Reducing Metabolic Rate 121
Cyanide and Nitrates in Mine Spoils 169
Hibernation by a Tropical Species 121
Concept 5.5 Review 123
Applications: Local Extinction of a Land Snail in an Urban Chapter 8 Social Relations 172
Heat Island 123 Concepts 172
8.1 Mate Choice versus Predation 174
Chapter 6 Water Relations 127 Mate Choice and Sexual Selection in Guppies 175
Concepts 127 Concept 8.1 Review 178
6.1 Water Availability 128 8.2 Mate Choice and Resource Provisioning 178
Water Content of Air 129 Concept 8.2 Review 181
Water Movement in Aquatic Environments 130 8.3 Nonrandom Mating in a Plant Population 181
Water Movement Between Soils and Plants 131 Concept 8.3 Review 183
Concept 6.1 Review 132 8.4 Sociality 183
6.2 Water Regulation on Land 133 Cooperative Breeders 184
Water Acquisition by Animals 134 Concept 8.4 Review 189
viii Contents

8.5 Eusociality 189 10.3 Patterns of Survival 224

Eusocial Species 189 Estimating Patterns of Survival 224
Evolution of Eusociality 191 High Survival Among the Young 224
Concept 8.5 Review 193 Constant Rates of Survival 226
Applications: Behavioral Ecology and Conservation 193 High Mortality Among the Young 227
Three Types of Survivorship Curves 227
Tinbergen’s Framework 193
Concept 10.3 Review 228
Environmental Enrichment and Development of
Behavior 193 10.4 Age Distribution 228
Contrasting Tree Populations 228
A Dynamic Population in a Variable Climate 229
Section III Concept 10.4 Review 230
10.5 Rates of Population Change 230
POPULATION ECOLOGY Estimating Rates for an Annual Plant 230
Estimating Rates When Generations Overlap 231
Chapter 9Population Distribution and Concept 10.5 Review 233
Abundance 196 Applications: Changes in Species Distributions in Response to
Climate Warming 233
Concepts 196
9.1 Distribution Limits 198
Kangaroo Distributions and Climate 198 Chapter 11 Population Growth 237
Distributions of Plants Along a Moisture-Temperature Concepts 237
Gradient 199
11.1 Geometric and Exponential Population Growth 238
Distributions of Barnacles Along an Intertidal Exposure
Geometric Growth 238
Gradient 200
Exponential Growth 240
Concept 9.1 Review 202
Exponential Growth in Nature 241
9.2 Patterns on Small Scales 202
Concept 11.1 Review 242
Scale, Distributions, and Mechanisms 202
11.2 Logistic Population Growth 242
Distributions of Tropical Bee Colonies 202
Concept 11.2 Review 245
Distributions of Desert Shrubs 204
Concept 9.2 Review 205 11.3 Limits to Population Growth 245
9.3 Patterns on Large Scales 205 Environment and Birth and Death Among Darwin’s
Finches 245
Bird Populations Across North America 206
Concept 11.3 Review 248
Plant Distributions Along Moisture Gradients 207
Concept 9.3 Review 208 Applications: The Human Population 248

9.4 Organism Size and Population Density 208 Distribution and Abundance 248
Population Dynamics 249
Animal Size and Population Density 208
Population Growth 250
Plant Size and Population Density 209
Concept 9.4 Review 210
Applications: Rarity and Vulnerability to Extinction 210 Chapter 12 Life Histories 254
Seven Forms of Rarity and One of Abundance 210 Concepts 254
12.1 Offspring Number versus Size 255
Chapter 10 Population Dynamics 215 Egg Size and Number in Fish 256
Seed Size and Number in Plants 258
Concepts 215 Seed Size and Seedling Performance 259
10.1 Dispersal 217 Concept 12.1 Review 261
Dispersal of Expanding Populations 217 12.2 Adult Survival and ­Reproductive Allocation 262
Range Changes in Response to Climate Change 218 Life History Variation Among Species 262
Dispersal in Response to Changing Food Supply 219 Life History Variation within Species 264
Dispersal in Rivers and Streams 220 Concept 12.2 Review 266
Concept 10.1 Review 221 12.3 Life History Classification 266
10.2 Metapopulations 221 r and K Selection 266
A Metapopulation of an Alpine Butterfly 222 Plant Life Histories 267
Dispersal Within a Metapopulation of Lesser Kestrels 223 Opportunistic, Equilibrium, and Periodic Life
Concept 10.2 Review 224 Histories 268
 Contents ix

Lifetime Reproductive Effort and Relative Offspring Size: Experimental Test of Food and Predation Impacts 306
Two Central Variables? 270 Population Cycles in Mathematical and Laboratory
Concept 12.3 Review 272 Models 307
Applications: Climate Change and Timing of Reproduction and Concept 14.2 Review 310
Migration 272 14.3 Refuges 310
Altered Plant Phenology 272 Refuges and Host Persistence in Laboratory and
Animal Phenology 273 ­Mathematical Models 310
Exploited Organisms and Their Wide Variety of

Section IV “Refuges” 312

Concept 14.3 Review 314
INTERACTIONS 14.4 Ratio-Dependent Models of ­Functional Response 314
Alternative Model for Trophic Ecology 314
Chapter 13
Species Interactions and Evidence for Ratio-Dependent Predation 315
Competition 277 Concept 14.4 Review 317
14.5 Complex Interactions 317
Concepts 277
Parasites and Pathogens That Manipulate Host Behavior 317
Competitive Interactions Are Diverse 279 The Entangling of Exploitation with Competition 320
13.1 Intraspecific Competition 280 Concept 14.5 Review 321
Intraspecific Competition Among Plants 280 Applications: The Value of Pest Control by Bats: A Case
Intraspecific Competition Among Planthoppers 281 Study 321
Interference Competition Among Terrestrial
Isopods 282
Concept 13.1 Review 282 Chapter 15 Mutualism 325
13.2 Competitive Exclusion and Niches 282 Concepts 325
The Feeding Niches of Darwin’s Finches 283 15.1 Plant Mutualisms 326
Competition for Caterpillars 284 Plant Performance and Mycorrhizal Fungi 327
Concept 13.2 Review 285 Ants and Swollen Thorn Acacias 330
13.3 Mathematical and Laboratory Models 285 A Temperate Plant Protection Mutualism 334
Modeling Interspecific Competition 285 Concept 15.1 Review 335
Laboratory Models of Competition 288 15.2 Coral Mutualisms 335
Concept 13.3 Review 289 Zooxanthellae and Corals 336
13.4 Competition and Niches 289 A Coral Protection Mutualism 336
Niches and Competition Among Plants 289 Concept 15.2 Review 338
Niche Overlap and Competition Between Barnacles 290 15.3 Evolution of Mutualism 338
Competition and the Niches of Small Rodents 291 Facultative Ant-Plant Protection Mutualisms 340
Character Displacement 293 Concept 15.3 Review 341
Evidence for Competition in Nature 295 Applications: Mutualism and Humans 341
Concept 13.4 Review 295 Guiding Behavior 341
Applications: Competition Between Native and Invasive
Species 296
Section V
Chapter 14 Exploitative Interactions: COMMUNITIES AND ECOSYSTEMS 345
Predation, Herbivory,
Parasitism, and Disease 299 Chapter 16 Species Abundance
and Diversity 345
Concepts 299
Concepts 345
14.1 Exploitation and Abundance 300
A Herbivorous Stream Insect and Its Algal Food 300 16.1 Species Abundance 347
Bats, Birds, and Herbivory in a Tropical Forest 301 The Lognormal Distribution 347
A Pathogenic Parasite, a Predator, and Its Prey 303 Concept 16.1 Review 348
Concept 14.1 Review 304 16.2 Species Diversity 348
14.2 Dynamics 304 A Quantitative Index of Species Diversity 348
Cycles of Abundance in Snowshoe Hares and Their Rank-Abundance Curves 350
Predators 304 Concept 16.2 Review 351
x Contents

16.3 Environmental Complexity 351 18.2 Patterns of Aquatic Primary Production 387
Forest Complexity and Bird Species Diversity 351 Patterns and Models 387
Niches, Heterogeneity, and the Diversity of Algae and Whole-Lake Experiments on Primary Production 388
Plants 352 Global Patterns of Marine Primary Production 388
The Niches of Algae and Terrestrial Plants 353 Concept 18.2 Review 389
Complexity in Plant Environments 354 18.3 Primary Producer Diversity 390
Soil and Topographic Heterogeneity 354 Terrestrial Plant Diversity and Primary Production 390
Nutrient Enrichment Can Reduce Environmental Algal Diversity and Aquatic Primary Production 391
Complexity 355
Concept 18.3 Review 391
Nitrogen Enrichment and Ectomycorrhizal Fungus
Diversity 356 18.4 Consumer Influences 392
Concept 16.3 Review 357 Piscivores, Planktivores, and Lake
Primary Production 392
16.4 Disturbance and Diversity 357
Grazing by Large Mammals and Primary Production on
The Nature and Sources of Disturbance 357 the Serengeti 394
The Intermediate Disturbance Hypothesis 357 Concept 18.4 Review 396
Disturbance and Diversity in the Intertidal Zone 358
18.5 Secondary Production 396
Disturbance and Diversity in Temperate Grasslands 358
A Trophic Dynamic View of Ecosystems 397
Concept 16.4 Review 360
Top-down Versus Bottom-up Controls on Secondary
Applications: Disturbance by Humans 360 Production 397
Urban Diversity 361 Linking Primary Production and Secondary
Production 398
Chapter 17 Species Interactions and Concept 18.5 Review 399
­Community Structure 365 Applications: Using Stable Isotope Analysis to Study Feeding
Habits 399
Concepts 365
Using Stable Isotopes to Identify Sources of Energy in a
17.1 Community Webs Salt Marsh 400
Strong Interactions and Food Web Structure 367
Concept 17.1 Review 368
17.2 Indirect Interactions 368
Chapter 19 Nutrient Cycling and
Retention 403
Indirect Commensalism 368
Apparent Competition 369 Concepts 403
Concept 17.2 Review 370 19.1 Nutrient Cycles 404
17.3 Keystone Species 371 The Phosphorus Cycle 405
Food Web Structure and Species Diversity 371 The Nitrogen Cycle 406
Experimental Removal of Sea Stars 373 The Carbon Cycle 407
Snail Effects on Algal Diversity 374 Concept 19.1 Review 408
Fish as Keystone Species in River Food Webs 376 19.2 Rates of Decomposition 408
Concept 17.3 Review 377 Decomposition in Two Mediterranean Woodland
17.4 Mutualistic Keystones 378 Ecosystems 408
A Cleaner Fish as a Keystone Species 378 Decomposition in Two Temperate Forest
Seed Dispersal Mutualists as Keystone Species 379 Ecosystems 409
Concept 17.4 Review 379 Decomposition in Aquatic Ecosystems 411
Applications: Human Modification of Food Webs 380 Concept 19.2 Review 412
Parasitoid Wasps: Apparent Competition and Biological 19.3 Organisms and Nutrients 412
Control 380 Nutrient Cycling in Streams and Lakes 412
Animals and Nutrient Cycling in Terrestrial
Chapter 18 Primary and Secondary Ecosystems 415
Plants and the Nutrient Dynamics of Ecosystems 416
Production 383 Concept 19.3 Review 417
Concepts 383 19.4 Disturbance and Nutrients 417
18.1 Patterns of Terrestrial Primary Production 385 Disturbance and Nutrient Loss from Forests 417
Actual Evapotranspiration and Terrestrial Primary Flooding and Nutrient Export by Streams 418
Production 385 Concept 19.4 Review 419
Soil Fertility and Terrestrial Primary Production 386 Applications: Altering Aquatic and Terrestrial
Concept 18.1 Review 387 Ecosystems 419
 Contents xi

Chapter 20 Succession and Stability 423 Chapter 22 Geographic Ecology 468

Concepts 423 Concepts 468
20.1 Community Changes During Succession 425 22.1 Area, Isolation, and Species Richness 470
Primary Succession at Glacier Bay 425 Island Area and Species Richness 470
Secondary Succession in Temperate Forests 427 Island Isolation and Species Richness 472
Succession in Rocky Intertidal Communities 427 Concept 22.1 Review 473
Succession in Stream Communities 428 22.2 The Equilibrium Model of Island Biogeography 473
Concept 20.1 Review 429 Species Turnover on Islands 474
20.2 Ecosystem Changes During Succession 429 Experimental Island Biogeography 475
Four Million Years of Ecosystem Change 429 Colonization of New Islands by Plants 476
Succession and Stream Ecosystem Properties 431 Manipulating Island Area 477
Concept 20.2 Review 432 Island Biogeography Update 478
20.3 Mechanisms of Succession 432 Concept 22.2 Review 478
Facilitation 433 22.3 Latitudinal Gradients in Species Richness 478
Tolerance 433 Latitudinal Gradient Hypotheses 478
Inhibition 433 Area and Latitudinal Gradients in Species Richness 480
Successional Mechanisms in the Rocky Intertidal Zone 434 Continental Area and Species Richness 481
Mechanisms in Old Field Succession 435 Concept 22.3 Review 482
Concept 20.3 Review 436 22.4 Historical and Regional Influences 482
20.4 Community and Ecosystem Stability 436 Exceptional Patterns of Diversity 482
Lessons from the Park Grass Experiment 437 Historical and Regional Explanations 483
Replicate Disturbances and Desert Stream Stability 438 Concept 22.4 Review 484
Concept 20.4 Review 440 Applications: Global Positioning Systems, Remote Sensing,
Applications: Ecological Succession Informing Ecological and Geographic Information Systems 485
Restoration 440 Global Positioning Systems 485
Applying Succession Concepts to Restoration 440 Remote Sensing 485
Geographic Information Systems 487

SECTION VI Chapter 23 Global Ecology 490

LARGE-SCALE ECOLOGY Concepts 490
The Atmospheric Envelope and the Greenhouse
Chapter 21 Landscape Ecology 445 Earth 491
Concepts 445 23.1 A Global System 493
The Historical Thread 493
21.1 Landscape Structure 447
El Niño and La Niña 494
The Structure of Six Landscapes in Ohio 447
El Niño Southern Oscillation and Marine
The Fractal Geometry of Landscapes 449
Populations 495
Concept 21.1 Review 450
El Niño and the Great Salt Lake 497
21.2 Landscape Processes 450 El Niño and Terrestrial Populations in Australia 498
Landscape Structure and the Dispersal of Mammals 451 Concept 23.1 Review 499
Habitat Patch Size and Isolation and the Density of 23.2 Human Activity and the Global Nitrogen Cycle 499
­Butterfly Populations 452
Concept 23.2 Review 500
Habitat Corridors and Movement of Organisms 453
Landscape Position and Lake Chemistry 454 23.3 Changes in Land Cover 500
Concept 21.2 Review 455 Deforestation 500
Concept 23.3 Review 504
21.3 Origins of Landscape Structure and Change 455
Geological Processes, Climate, and Landscape 23.4 Human Influence on ­Atmospheric Composition 504
Structure 456 Depletion and Recovery of the Ozone Layer 507
Organisms and Landscape Structure 458 Concept 23.4 Review 508
Fire and the Structure of a Mediterranean Landscape 462 Applications: Impacts of Global Climate Change 508
Concept 21.3 Review 463 Shifts in Biodiversity and Widespread Extinction of
Applications: Landscape Approaches to Mitigating Urban Heat Species 509
Islands 463 Human Impacts of Climate Change 509
xii Contents

Appendix A Investigating the Evidence

16: Estimating the Number of Species in Communities 531
1: The Scientific Method—Questions and Hypotheses 514 17: Using Confidence Intervals to Compare
2: Determining the Sample Mean 515 Populations 532
3: Determining the Sample Median 516 18: Comparing Two Populations with the t-Test 533
4: Variation in Data 517 19: Assumptions for Statistical Tests 534
5: Laboratory Experiments 518 20: Variation Around the Median 535
6: Sample Size 519 21: Comparison of Two Samples Using a Rank Sum
7: Scatter Plots and the Relationship Between Test 537
Variables 520 22: Sample Size Revisited 538
8: Estimating Heritability Using Regression 23: Discovering What’s Been Discovered 539
Analysis 521
Appendix B Statistical Tables 541
9: Clumped, Random, and Regular Distributions 522
10: Hypotheses and Statistical Significance 523 Appendix C Abbreviations Used in This Text 545
11: Frequency of Alternative Phenotypes in a Appendix D Global Biomes 547
Population 524
12: A Statistical Test for Distribution Pattern 526 Glossary 548
13: Field Experiments 527 References 558
14: Standard Error of the Mean 528
15: Confidence Intervals 530 Index 571

Design elements: Anna A. Sher

 Preface
This book was written for students taking their first under- example of an ecological process. All attempt to engage stu-
graduate course in ecology. We have assumed that students dents and draw them into the discussion that 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, which are listed
biology, which included introductions to evolution, physiol-
at the beginning of each chapter to alert the student to
ogy, and biological diversity.
the major topics to follow and to provide a place where the
student can find a list of the important points covered in
Organization of the Book each chapter. The sections in which concepts are discussed
An evolutionary perspective forms the foundation of the focus on published studies and, wherever possible, the sci-
entire textbook, as it is needed to support understanding of entists who did the research are introduced. This case-study
major concepts. The textbook begins with a brief introduction approach supports the concepts with evidence, and intro-
to the nature and history of the discipline of ecology, followed duces students to the methods and people that have created
by section I, which includes two chapters on earth’s biomes— the discipline of ecology. Each concept discussion ends
life on land and life in water—followed by a chapter on popu- with a series of concept review questions to help students
lation genetics and natural selection. Sections II through VI test their knowledge and to reinforce key points made in
build a hierarchical perspective through the traditional sub- the discussion.
disciplines of ecology: section II concerns adaptations to
the environment; section III focuses on population ecology;
section IV presents the ecology of interactions; section V
summarizes community and ecosystem ecology; and finally,
First Pages
section VI discusses large-scale ecology, including chapters
on landscape, geographic, and global ecology. These topics
were first introduced in section I within its discussion of
SEC TIO N
II Adaptations
to the Environ

Chapter
ment

the biomes. In summary, the book begins with an overview

of the biosphere, considers portions of the whole in the

5
middle chapters, and ends with another perspective of the
entire planet in the concluding chapter. The features of this
textbook were carefully planned to enhance the students’
comprehension of the broad discipline of ecology.
Temperature
Features Designed with the Student in Relations
Mind ©Digital Vision
/Getty Images
RF
Japanese maca
ques
body heat in the , Macaca fuscata, huddle toget
All chapters are based on a distinctive learning system, fea- mids
temperature, using t of driving snow. The capa
tions, enables these
behavioral, anat
her, conserving
city to regulate
omical, and phys
their
body
iological adapta-
Japan, site of the monkeys to live through the 5.5 Many orga
turing the following key components: 1998 Winter Olym
pics.
cold winters in
Nagano, nisms surv
temperatures
stage. 120
ive extreme
by entering a
resting
Concept 5.5 Rev
CHAPTER CO iew 123
Student Learning Outcomes: Educators are being asked NCEPTS Applications:
Local Extinct
an Urban Hea ion of a Land
Snail in
increasingly to develop concrete student learning outcomes 5.1 Macroclima
landscape to
te inte racts with the
produce microcl local
Summary 124
Key Terms
t Island 123

variation in tem ima

perature. 102 tic 125
for courses across the curriculum. In response to this need Concept 5.1 Rev
iew 105
Review Questio
ns 125
5.2 Adapting
and to help focus student progress through the content, all to one
conditions gen set of environmental
a population’
erally reduce
s
sections of each chapter in the ninth edition begin with a list
s fitness in oth
environments. er
105 LEARNING OU
Concept 5.2 Rev TCOMES
iew 106 After studying this
of detailed student learning outcomes. 5.3 Most species
narrow range
perform best
in a fairly
5.1
section you shou
Distinguish betw
een temperature
ld be able to do
the
following:
of temperatu 5.2 Explain the ecol and heat.
Concept 5.3 Rev res. 107 ogical significa
iew 110 tal temperature nce of environm
en-
Introduction: The introduction to each chapter presents
s.
5.4 Many orga

T
nisms
ways to compen have evolved he thermometer
the student with the flavor of the subject and important in environmen
regulating bod
sate for variatio
tal temperatu
y temperature.
re by
ns appear in the scie
suring and repo
was one of the
ntific tool kit, and
first instruments
we have been mea
to
what do thermom rting temperatures ever sinc -
background information. Some introductions include Concept 5.4 Rev 110 e. However,
iew 120 eters actually
measure of the quantify? Tem
average kinetic perature is a
the molecules, energy, or ener
in a mass of a gy of motion, of
historical events related to the subject; others present an substance. For
example, water

101

moL22201_ch
05_101-126.indd
101

02/16/21 10:33
AM

xiii
 xiv Preface First Pages

Illustrations: A great deal of effort has been put into In the mid-190
0’s, darker pep
Chapter 7
Energy and Nu
trient Relatio
ns
the development of illustrations, both photographs moths were less
soot-covered tree
s and so had a
pered
visible to predator
s on
fitness ... whereas by the
161
advantage over
and line art. The goal has been to create more-effec- lighter moths... 1990’s, air qual
improved, and ity
lighter moths beca
better camoufl me the
aged morph...

tive pedagogical tools through skillful design and

use of color, and to rearrange the traditional presen-
tation of information in figures and captions. Much
explanatory material is located within the illustra-
tions, providing students with key information where
they need it most. The approach also ­provides an
ongoing tutorial on graph interpretation, a skill with
which many introductory students need practice.
1959
7%
Detailed Explanations of ­Mathematics: The math- 1996
ematical aspects of ecology commonly challenge Dark morph abun
dance 8%
93% Light morph abun
many students taking their first ecology course. dance

This text carefully explains all mathematical expres- 92%

sions that arise to help students overcome these Figure 7.16

... such that the
the morphs reve
relative abundan
ce of
Birds and othe rsed; light beca
challenges. In some cases, mathematical expressions extent to which
wake of environ
they are camouf
men
r predators exer
laged. When tree
t strong selectiv
s wer
e pressure on pop
ulations of pep
common than
dark.
me more

of the lighter mor tal regulations that reduced air e covered with soot due to indu pered moths in
are dissected in illustrations designed to comple- examples of natu
Bill Coster IN/A
ph. Frequency
data shown here
ral selection, alth
oug h rece
pollution, the bark
were collected
37 year
of trees became
s apa
strialization, dark
light-colored agai
Europe and else
er morphs were
more
where, based on
n, resulting in dram abundant, whereas in the
the
lamy Stock Phot nt research sugg rt in West Kirb
ment their presentation in the associated narrative. o/H Lansdown/Al ests that other y, Wirral. This atic shifts in the
amy Stock Phot evolutionary forc remains one of frequency
o/Frank Hecker/A es the mos
First Pages lamy Stock Phot
o
also likely played a
role (data from
t commonly-used
Grant et al. 199
8).
Because predat
ors must catch
often select pre and subdue the
y by size, a beh ir
selective predat avior that ecolog prey, they diff icult to find
ion. Because of ists call size- or catch. As we
significantly cor this behavior, pre size-selective pre shall see later
related with pre y size is often dation may also in chapter 7,
A visualization of a population bottle neck, using data from
solitar y predat dator size, esp Size of the predat have an energe
Chapter 10 Population Dynamics ors. One such ecially among or can affect pre tic basis.
mountain lion solitar y predat Shannon Murph y selection in oth
225 , Puma concolo or, the puma, y, Danny Lewis, er ways.
published research.
Yukon to the r, ranges from or impacts of size and Gina Wim
tip of South Am the Canadian by asking if the p studied the
changes substa erica (fig. 7.18 ralis) in salt ma diets of wolf spid
To allow comparisons to ntially along this ). Puma size rshes changed as ers (Pardosa litto
other tin Iriarte and latitudinal gra Older spiders are they aged (Murp -
studies, number of Dall sheep Subtracting number of his colleagues dient. Augus- as much as 30 tim hy et al. 2020).
deaths from number alive increase in size (1990) found spiders; howeve es larger than new
surviving and dying withi , the average size that as r, in theory sma ly hatched
n each the beginning of each year
at (fig. 7.19). Ma of their prey also pumas same diet becaus ll and large spid
e they digest the ers could have the
year of life is converted to mmals make up increases
gives the number alive at and large mamm over 90% of the have to be larger ir prey externally
number per 1,000 births. the als, especially puma’s diet, than their prey. and so don’t
beginning of the next year. nor thern par t deer, are its ma (see chapter 6) Using nitrogen
of its range in in prey in the of field-collected isotope analysis
are larger. In the North Americ mine that diet did spiders, they wer
tro a, wh ere cha nge: young spid e able to
Number of mainly on mediu pics where pumas are smalle pumas tovores, which occ ers primarily hun deter-
m and small pre r, they feed ur on the soil sur ted detri-
should differe y, especially rod the larger, older face under the tha
Age (years) survivors Number of deaths nt-sized pumas ents. Why wolf spiders wer tch, whereas
One reason is feed on differe open. However, e hunting herbivo
at beginning during year that large pre nt-sized prey? a few small spid res out in the
and may even y ma to large spiders, ers had isotope
of year injure the predat y be diff icult to subdue confirming that
there was nothin
signatures similar
or, while small that kept them g physiological
0–1 1,000 prey may be from eating the
199 her colleagues sam e herbivore diet
1–2 1,000–199 concluded that
the mechanism
. Murphy and Chapter 18 Primary and Secon
801 12 for the differen
2–3 789 801–12 13 ce
3–4 By reducing planktivorous
776 789–13 12 fish
4–5 populations, piscivores indir
ectly and introduced them to
764 30 moL22201_ch0
7_149-171.indd increase populations of large th
5–6 734
161
unmanipulated as a con
6–7 etc. 46 zooplankton and indirectly
reduce
trol
688 48 biomass of phytoplankton. The responses of the
7–8 640 69 Lake food web 02/22/21 02:57
PM tal manipulations suppor
8–9 571 t
132 (see fig. 18.14). Reducing
9–10 439 187 the
10–11 to reduced rates of primary
252 156 p
11–12 96 tivorous minnows, the pred
12–13
90 ace
6 Piscivores more numerous. Chaobor
13–14
3 us
3 3 bivorous zooplankton, and
14–15 0 th
blage shifted in dominan
Planktivorous fish ce fr
Planktivorous presence of abundant, larg
li, a mountain sheep of far
Plotting age on the x-axis e
northern and number of survivors invertebrates plankton biomass and rate
e of the classic studies of
survivor- on the y-axis creates a
of
Adding planktivorous
survivorship curve. m
production

ecological response.
Incre
Dall sheep surviving their Large herbivorous Small herbivorous population led to incr
at that age. For example, of life have a high probability
first year
zooplankton eased
the of zooplankton However, though the rese
e that died before the age surviving to about age 9. arch
ary

of planktivorous fish in this

n that generally dies duri expe
Top-down influences on prim

ng 1,000
ssumption is not likely to unintended way. Despite
be the
l that emerges probably gives a few bass remained. So,
Sheep 10 years by
the population, particula old and older are minnows, they basically
rie’s.
rly fed th
easier prey for food supply combined
Number of survivors

e survival patterns for

100 wolves and die at Large phytoplankton with
Dall a high rate. Small phytoplankton induced a strong numeric
skulls. The upper portion Survivorship curves are al re
of (see chapter 10). The man
that Murie constructed. The plotted using a log ipu
eep, the second column lists scale on the y-axis.10 tive rate of the remaining
large
class, and the third colu 10 an abundance of young
mn largemo
class. Notice that although on zooplankton.
e numbers in the table The lake ecosystem resp
are on
ividuals. This adjustment of planktivorous fish (you
is
er populations. 1 ng la
14 also shows how to tran
at the outset of the experim
s-
0 2 4 6 8 10
Nutrients ent.
bers of survivors. Plotting 12 14 Figure 18.12 The trop decreased sharply, the aver
Age (years) hic cascade hypothesis, a
“cascading” indirect inter result of age si
s against age produces the Figure 10.14 Dall sheep actions. Lines indicate both decreased, and phytopl
: from life table to survivorsh indirect (dashed) relations direct (solid) and ankto
wer portion of the figur
e.
(data from Murie 1944). ip curve hips. production increased.
s of life and death within
opulation of Dall sheep,
overall pattern of survival
y rates are higher: during and mor tality among Dall
Helps
between 9 and students
13 years.
much likeworkthat for awith
varietyand of otheinterpret
r large vertebrate quantitative
sheep is informa- Provides a visual representation of a hypothesis involving ael
The trophic cascade mod
predicts that manipulating
ing red deer, Cervus elaph s, includ-
tion, involving
he aged are higher in this
Odocoileuconverting
s hem ionu numerical
us, Colu mbia
s columbianus, East Afri
n information
black-tailed deer, into a graph. set of complex ecological interactions. pisci vore biomass will lead to
iddle years is lower. The can buffalo, Syn- changes in biomass and
cerus caffer, and humans.
The key characteristics production of planktivores,
of survival
herbivores, and phytopla
nkton.

Planktivores
Discovering Diverse Content Through
Random Scribd Documents
Fauna from the Caves of Mentone 373
” Bone-caves of Sicily 376
List of Animals from the Middle Pleistocene 415
” ” ” Early Pleistocene 418
” Pleistocene Mammalia 420, 422
” Characteristic Animals of the Pleistocene Period 423
” ” ” ” Pleiocene Period 424
 ADDITIONS AND CORRECTIONS.
Page 1, line 7, for “Cythæron” read “Cithæron.”
Page 8, line 4, for “that” read “who.”
Page 17, line 5, for “Seine” read “Somme.”
Page 60, lines 29, 30, for “non-ossiferous” read “no ossiferous.”
Page 82, fig. 19, for “A, B, Albert, C, Victoria” read “A, B, Victoria,
C, Albert.”

Page 95, fig. 25.—This design is to be seen in the chalice

discovered in 1868, in a rath at Ardagh, Limerick, and described by
the Earl of Dunraven (Trans. Royal Irish Acad. xxiv. Antiquities). The
chalice is made of gold, silver, bronze, brass, copper, and lead, and
from the identity of its inscription and ornament with those of Irish
MSS. of ascertained age, may be referred to a date ranging from the
5th to the 9th centuries. It is also adorned with squares of blue and
red enamel of the same kind as that of the brooches from the
Victoria Cave, figured in the coloured plate. The same design is also
presented by the “bronze head-ring” found in 1747 at Stitchel, in
Roxburgh, (Wilson “Prehistoric Annals of Scotland,” ii. 146) as well as
by one of the silver articles known as “The Norrie Law Relics,” found
in a tumulus on the shore of the Bay of Largo, Firth of Forth. Of the
coins found at the same place, the latest, belonging to Tiberius
Constantine (d. 682), fixes the date as not earlier than the 7th
century. Some of the sculptured stones of Scotland, such as the
Dunnichen stone, are ornamented also in the same style, and,
according to Professor Wilson, belong to “the transition period from
the 4th to the 8th centuries, when pagan and Christian rites were
obscurely mingled,” (ii. 259). In Scotland, therefore, as well as
Ireland, this style of ornamentation is of the same age,
corresponding in the main with that of Brit-Welsh articles in the
Victoria Cave, proved by the associated coins to be later than the
4th century.
Page 120, line 4.—These teeth are considered by Dr. Leith
Adams to belong to Elephas antiquus, which has been discovered in
other places in Yorkshire. They may possibly belong to that animal;
but they may, with equal justice, be identified with the wide-plated
variety of the teeth of the Mammoth. The great variation in the
width of the component plates of the fossil teeth of Mammoth
observable in the large series from Crayford and the caves of the
Mendip Hills, and in those in the magnificent Museum of Lyons,
causes me to hesitate in considering them to belong to the rarer
species.
Page 130, line 2.—This has been verified while these sheets were
passing through the press by the discovery of Brit-Welsh articles in a
cave in Kirkcudbrightshire by Messrs. A. R. Hunt and A. J. Corrie,
among which are bone fasteners similar in outline to that from the
Victoria Cave (Fig. 23).
Page 190.—In using this classification of crania, I have purposely
attached higher value to the two extremes of skull form, or the long
and the broad, than to the intermediate oval forms, which cannot be
viewed as distinctive of race, because they may be the results either
of the intermarriage of a long-headed with a short-headed people,
or of variation from the type of one or other of them.
Page 196, heading, for “Dolicho-cepha” read “Dolicho-cephali.”
Page 201, heading, dele “A”.
Page 213, note 2.—The “tête annulaire,” or annular depression,
is also visible on some of the broad as well as the long skulls from a
“Merovingian” cemetery at Chelles in the same collection. The
association in this cemetery of the two skull-forms is probably due to
the Merovingians being the masters, and the Celts the servants, and
the conquerors and the vanquished being buried in the same spot.
Page 220, line 24, for “Volscæ” read “Volcæ.”
Page 223, line 25, for “east” read “west.”
Page 228, line 3, dele “that.”
Page 229, line 3, for “set foot” read “settled.” The statement in
the text is too strong. The conquest of Gaul by the Huns under Attila
was averted by his defeat in the famous battle of Chalons.
Page 275, line 21, for “are” read “is.”
Page 279.—Since this was written a new ossiferous deposit has
been found in a fissure at Lothorsdale, near Skipton, from which the
remains of the Elephas antiquus and Hippopotamus amphibius have
been obtained.
Page 284.—The ossiferous fissure at Windy Knoll, near Castleton,
recently explored by Messrs. Tym, Pennington, Plant, Walker and
others, has added several animals to the pleistocene fauna of that
district—the bison, roe, reindeer, bear, wolf, fox, and hyæna, the first
of these species being remarkably abundant, and of all ages. The
remains were probably introduced by a stream from a higher level.
Page 337, note 2, line 2, for “the Revue” and “les Matériaux”
read “in the Revue” and “in the Matériaux.”
Page 337, note 5, for “Aquitainicæ” read “Aquitanicæ.”
Page 347, line 6, for “mind” read “minds.”
Page 356, line 15, for “Port” read “Fort.”
Page 361.—Mr. Ayshford Sanford adds the Felis Caffer to the list
from Bleadon, and the Gulo borealis to that of the animals from
Kent’s Hole.
Page 386, line 10, dele inverted commas.
Page 386, line 17, for “or from 1,000 to 2,000 feet lower than
the glacial covering” read “thus differing by a line of from 1,000 to
2,000 feet from the glacial covering” (Palgrave).
CAVE-HUNTING.
 CHAPTER I.
INTRODUCTION.
Legends and Superstitions connected with Caves.—The Physical Division
of the Subject.—The Biological.—The Inhabitants of Caves.—Men and
Animals.—Ethnological, Archæological, and Geographical Bearings.—
The three Classes of Bone-Caves: Historic, Prehistoric, Pleistocene.—
History of Cave Exploration in Europe: Germany, Great Britain, France,
Belgium, Southern Europe.

Caves have excited the awe and wonder of mankind in all ages, and
have figured largely in many legends and superstitions. In the
Roman Mythology, they were the abode of the Sibyls, and of the
nymphs, and in Greece they were the places where Pan, Bacchus,
Pluto, and the Moon were worshipped, and where the oracles were
delivered, as at Delphi, Corinth, and Mount Cithæron; in Persia they
were connected with the obscure worship of Mithras. Their names,
in many cases, are survivals of the superstitious ideas of antiquity. In
France and Germany they are frequently termed “Fairy, Dragons’, or
Devils’ Caves,” and, according to M. Desnoyers, they are mentioned
in the invocation of certain canonized anchorites, who dwelt in them
after having dispossessed and destroyed the dragons and serpents,
the pagan superstition appearing in a Christian dress.
In the Middle Ages they were looked upon as the dwellings of
evil spirits, into the unfathomable abysses of which the intruder was
lured to his own destruction. Long after the fairies and little men had
forsaken the forests and glens of Northern Germany, they dwelt in
their palaces deep in the hearts of the mountains,—in “the dwarf
holes,” as they were called—whence they came, from time to time,
into the upper air. Near Elbingrode, for example, in the Hartz, the
legend was current in the middle of the last century, that when a
wedding-dinner was being prepared the near relations of the bride
and bridegroom went to the caves, and asked the dwarfs for copper
and brass kettles, pewter dishes and plates, and other kitchen
1
utensils. “Then they retired a little, and when they came back,
found everything they desired set ready for them at the mouth of
the cave. When the wedding was over they returned what they had
borrowed, and in token of gratitude, offered some meat to their
benefactors.” Allusions, such as this, to dwarfs, according to
Professor Nilsson, point back to the remote time when a small
primeval race, inhabiting Northern Germany, was driven by invaders
to take refuge in caverns,—a view that derives support from the fact
that in Scandinavia the tall Northmen were accustomed to consider
the smaller Lapps and Finns as dwarfs, and to invest them with
magic power, just as in Palestine the smaller invading peoples
considered their tall enemies giants. The cave of Bauman’s hole, also
in the Hartz district, was said, in the middle of the last century, to
have been haunted by divers apparitions, and to contain a treasure
guarded by black mastiffs; and in Burrington Combe, in
Somersetshire, some twenty years ago, a cave was dug out by a
working man, under the impression that it contained gold. The hills
of Granada are still believed, by the Moorish children, to contain the
great Boabdil and his sleeping host, who will awake when an
adventurous mortal invades their repose, and will issue forth to
restore the glory of the Moorish kings.
It is, indeed, no wonder that legends and poetical fancies such
as these should cluster round caves, for the gloom of their recesses,
and the shrill drip of the water from the roof, or the roar of the
subterranean water-falls echoing through the passages, and the
white bosses of stalagmite looming like statues through the
darkness, offer ample materials for the use of a vivid imagination.
The fact that often their length was unknown, naturally led to the
inference that they were passages into another world. And this is
equally true of the story of Boabdil, of that of the Purgatory of St.
Patrick, in the north of Ireland, and of the course of the river Styx,
which sinks into the rocks and flows through a series of caverns that
are the dark entrance-halls of Hades. The same idea is evident in the
remarkable story, related by Ælian (Lib. xvi. 16). “Among the Indians
of Areia there is an abyss sacred to Pluto, and beneath it vast
galleries, and hidden passages and depths, that have never been
fathomed. How these are formed the Indians tell not, nor shall I
attempt to relate. The Indians drive thither (every year) more than
3,000 different animals—sheep, goats, oxen, and horses—and each
acting either from dread of the dreadful abyss, or to avert an evil
omen in proportion to his means, seeks his own and his family’s
safety by causing the animals to tumble in; and these, neither bound
with chains nor driven, of their own accord finish their journey as if
led on by some charm; and after they have come to the mouth of
the abyss they willingly leap down, and are never more seen by
mortal eyes. The lowing, however, of the cattle, the bleating of the
sheep and of the goats, and the whinnying of the horses are heard
above ground, and if anyone listen at the mouth, he will hear sounds
of this kind lasting for a long time. Nor do they ever cease, because
beasts are driven thither every day. But whether the sound is made
by those recently driven in, or by some of those driven in some time
before, I do not express an opinion.” The Roman Catholic Church
took advantage of this feeling of superstitious awe, as late as the
Middle Ages. At the time of the Reformation it was believed that a
cave at Bishofferode would prove the death of some person in the
course of the year, unless a public yearly atonement were made.
Accordingly a priest came, on a certain day, to the chapel on the hill
opposite, whence he passed in solemn procession to the cave, “and
let down into it a crucifix, which he pulled up again, and took this
occasion to remind them of hell, and to avoid the punishment due to
their sins.”
The beauty of the interiors of some of the caves could not fail to
give rise to more graceful fancies than these. The fantastic shapes of
the dripstone, with which they are adorned, now resembling Gothic
pillars supporting a crystalline arcade, or jutting out in little spires
and minarets, and very generally covering the floor with a marble-
like pavement, and in some cases lining the pools of water with a
fretwork of crystals that shine like the facets of a diamond, were
fitting ornaments for the houses of unearthly beings, such as fairies.

The Physical Division of the Subject.

It is by no means my intention in this work to give a history of
legends such as these, but to take my readers with me into some of
the more important and more beautiful caves in this country. The
exploration of the chambers and passages of which they are
composed, the fording of the subterranean streams by which they
are frequently traversed, or the descent into deep chasms which
open in their floors, have the peculiar charm of mountaineering, not
without a certain pleasurable amount of risk. But to physicist and
geologist they offer far more than this. They give an insight into the
wonderful chemistry by which changes are being wrought, at the
present time, in the solid rock. Nor are the conclusions to which we
are led by the investigation of these chemical changes merely
confined to the interior of caves. They enable us to understand how
some of the most beautiful scenery in Europe has been formed, and
to realize the mode by which all precipices and gorges have been
carved out of the calcareous rock. In the next chapter we shall see
why it is that the combination of hill and valley, ravine and precipice,
present the same general features in all limestone districts—why, for
instance, the ravines of Palestine are the same as those of Greece,
and both are identical with those in Yorkshire. The origin and the
history of caves will be examined, as well as their relation to the
general physical geography of the calcareous strata. All these
subjects are comprehended in the first or the physical division of
cave-hunting.
 The Biological Division.
We must now proceed to the definition of the scope and object
of the second, or Biological, division of the subject.
Caves have been used by man, and the domestic animals living
under his protection, from the earliest times recorded by history
down to the present day. Those penetrating the rugged precipices of
Palestine, we read in the Old Testament, served both for habitation
and for burial, and, from the notices which are scattered through the
early Greek writers, we may conclude that those of Greece were
used for dwelling-places. The story of the Cyclops proves that they
were also used as folds for goats. The name of Troglodytes, given to
many peoples of the most remote antiquity, implies that there was a
time in the history of mankind when Pliny’s statement “specus erat
pro domibus” was strictly true (“Hist. Nat.” I. v. c. 56). The caves of
Africa have been places of retreat from the remotest antiquity down
to the French conquest of Algeria, and in 1845 several hundred
Arabs were suffocated in those of Dahra by the smoke of a fire
kindled at the entrance by Marshal (then Colonel) Pelissier. Dr.
Livingstone alludes in his recent letters to the vast caves of Central
Africa, which offer refuge to whole tribes with their cattle and
household stuff. In France, according to M. Desnoyers, there are at
the present time whole villages, including the church, to be found in
the rock, which are merely caves modified, extended, and altered by
the hand of man. The caves of the Dordogne were inhabited in the
middle ages. Floras writes that the Aquitani, “callidum genus in
2
speluncas se recipiebant, Cæsar jussit includi,” and the same caves
afforded shelter to the inhabitants of the same region in the wars of
King Pepin against the last Duke of Aquitaine. In this country a small
cave in Cheddar Pass was occupied till within the last few years. The
caves in the northern counties are stated by Gildas to have offered a
refuge to the Brit-Welsh inhabitants of Britain during the raids of the
Picts and Scots; and in the year 1745 those of Yorkshire were turned
to the same purpose during the invasion of the Pretender. We might
reasonably expect to find in caves turned to these uses objects left
behind, which would tell us something of the manners and customs
of their possessors, and light up the catalogue of battles and
intrigues of which history generally consists. The results obtained
from the Brit-Welsh group of caves, treated in the third chapter,
show that this hitherto neglected branch of the inquiry is not without
value to the historian.
Caves containing remains of this kind may be conveniently
termed historic, because they may be brought into relation with
history. It must, however, be carefully remarked that the term does
not relate to history in general, but to that in particular of each
country which happens to be under investigation. The
misapprehension of this has caused great confusion, and many
mistakes in archæological classification and reasoning.
Again, our experience of the habits of rude and uncivilized
peoples would naturally lead us to look to caves, as the places in
which we should be likely to meet with the remains of the men who
lived in Europe before the dawn of history. Such remains we do find
that, placed side by side with others from the tombs and dwellings,
enable us to discover some, at least, of the races who lived in
Europe in long-forgotten times, and to ascertain roughly the
sequence of events in the remote past, far away from the historical
border. It may, indeed, seem a hopeless quest to recover what has
been buried in oblivion so long, and it is successful merely through
the careful comparison of the human skeletons in the caves and
tombs of Britain, France, and Spain, with those of existing races, and
of the implements and weapons with those which are now used
among savage tribes. By this means we shall see that there are good
grounds for extending the range of the Iberian people over a
considerable area in Europe, and for the belief that the Eskimos once
lived as far south as Auvergne. In discussing both these problems it
will be impossible to shut our eyes to the continuity that exists
between geology, archæology, biology, and history—sciences which
at first sight appear isolated from each other.
 The bones of the domestic animals in the caves will necessarily
lead to the further examination of the appearance and
disappearance of breeds under the care of man. And this
complicated question has an important bearing not merely on the
ethnology, but also on the history, of some of the European peoples.
It must be admitted, however, that this branch of the subject is, as
yet, known merely in outline, and we can only hope to ascertain a
few facts which may form a basis for future investigation.
From another point of view the contents of caves are peculiarly
valuable. They have been used as places of shelter, not merely by
man, but by wild animals, from the time they first became accessible
to the present day. In the same way, therefore, as now they contain,
in their superficial layers, the bones of sheep, oxen, and horses,
foxes, rabbits, and badgers, so in their deeper strata lie buried the
remains of the animals which were living in Europe long before the
historic times. In other words, they enable us to make out the
groups of animals inhabiting the neighbouring districts, and which in
many cases have either forsaken their original abodes or have
become extinct. And since those which are extinct, or which have
migrated, could not have lived where their remains are found under
the present conditions of life, an inquiry into their history leads us
into the general question of the ancient European climate and
geography. It is obvious, for example, that the spotted hyæna,
which formerly inhabited the caves of Sicily, could not have crossed
over to that island after it was separated from Africa and Italy; and it
would be impossible for the musk-sheep, the most arctic of the
herbivora, to live as far south as Auvergne under the present
climatal conditions. The presence, therefore, of these animals in
these districts is proof in the one case of a geographical, and in the
other of a climatal, change.
The discussion of all these questions is comprehended under the
second, or biological, division of cave-hunting, which may be defined
as an inquiry into the remains of man and animals found in the
caves, and into the conditions under which they lived in Europe.
 The three Classes of Bone-caves.
In the biological branch of the subject the caves will be treated
first which are comprehended within the limits of history; then we
shall pass on to the investigation of Prehistoric caves, or those which
have been inhabited in the interval that separates history from the
remote geological era, which is characterized by the existence of the
extinct mammalia in Europe. And, lastly, those will be examined
which have furnished the remains of the extinct animals, and which
are termed by the geologists Pleistocene, from the fact that a larger
percentage of existing species were then living than in the preceding
Pleio-, Meio-, and Eocene periods. The equivalent terms
“Quaternary,” used by many French geologists, and the “Post-
pleiocene division of the Post-tertiary Formation,” used by Sir Charles
Lyell, are not adopted in this work, because they imply a break in
the continuity of life, which does not exist. “Pleistocene” was
3
invented and subsequently discarded by Sir C. Lyell, and is at
present used by many eminent writers, such as Forbes, Phillips,
Gervais, and others. The ossiferous caves will therefore be divided
into the Historic, Prehistoric, and Pleistocene groups. And it will be
more convenient to work backwards in time from the basis offered
by history, than to begin with the Pleistocene, or oldest division, and
bring the narrative down to the present day.
This classification, founded in part on the principle of change in
the animal world, and partly on the basis offered by history,
coincides, only in part, with that of the archæologists based on the
remains of man’s handiwork. The Pleistocene age is the equivalent
of the Palæolithic, or that of rude unpolished stone; the Prehistoric
represents the ages of polished stone, bronze, and iron in part, or
those stages in human progress when the use of these materials
became general for the purposes of every-day life; while the Historic
covers merely the later portion of that of iron.
 History of Cave-Exploration in Europe.
Germany.—The rest of this chapter must be devoted to an
outline of the history of cave-exploration during the last two
centuries. The dread of the supernatural, which preserved the
European caves from disturbance, was destroyed in the sixteenth
and seventeenth centuries by the search after “ebur fossile,” or
unicorn’s horn, which ranked high in the materia medica of those
days as a specific for many diseases, and which was obtained, in
great abundance, in the caverns of the Hartz, and in those of
Hungary and Franconia. As the true nature of the drug gradually
revealed itself, the German caves became famous for the remains of
the lions, hyænas, fossil elephants, and other strange animals, which
had been used for medicine. We owe the first philosophical
4
discussion on the point to Dr. Gesner, who, although he maintained
that the fossil unicorn consisted, in some cases, of elephant’s teeth
and tusks, and in others of its fossil bones, did not altogether give
up the idea of its medicinal value. It is a singular fact, that fossil
remains of a similar kind are, at the present time, used by the
5
Chinese for the same purpose, and sold in their druggists’ shops.
The cave which was most famous at the end of the seventeenth
century was that of Bauman’s Hole, in the Hartz, in the district of
Blankenbourg. It is noticed in the Philosophical Transactions for the
6
year 1662, and was subsequently described by Dr. Behrens,
Leibnitz, De Luc, and Cuvier, along with others in the
neighbourhood. Those of Hungary come next in point of discovery,
the first notice of them being due to Patterson Hayne in 1672. They
penetrate the southern slopes of the Carpathian ranges, and are
known by the name of dragons’ caves, because the bones which
they contain had been considered from time immemorial to belong
to those animals by the country people. These remains were
7
identified by Baron Cuvier as belonging to the cave-bear.
 It was not, however, until the close of the eighteenth century
that the exploring of caves was carried on systematically, or their
contents examined with any scientific precision. The caves of
Franconia, in the neighbourhood of Muggendorf, were described by
Esper in 1774, by Rosenmuller in 1804, and six years later by Dr.
Goldfuss. The most important was that of Gailenreuth, both from the
vast quantity of remains which it was proved to contain, and the
investigations to which it led. The bones of the hyæna, lion, wolf,
fox, glutton, and red deer were identified by Baron Cuvier; while
some of the skulls which Dr. Goldfuss obtained have been recently
proved, by Professor Busk, to belong to the grizzly bear. They were
associated with the bones of the reindeer, horse and bison.
Rosenmuller was of opinion that the cave had been inhabited by
bears for a long series of generations; and he thus realized that
these remains proved that the animals found in the cave had once
lived in that district, and had not been swept from the tropics by the
deluge. The interest in these discoveries was at its height in the year
1816, when Dr. Buckland visited the cave, and acquired that
knowledge of cave-exploring which he was subsequently to use with
8
such good effect in this country. From this time down to the present
day, no new fact of importance has been added to our knowledge of
caves by explorations in Germany.
Great Britain.—The first bone-cave systematically explored in this
9
country was that discovered by Mr. Whidbey, in the Devonian
limestone at Oreston, near Plymouth, in 1816; and the remains
obtained from it were identified by Sir Everard Home as implying the
existence of the rhinoceros in that region. This discovery followed
close upon the researches in Gailenreuth, and was due in some
degree to the request which Sir Joseph Banks made, that Mr.
Whidbey, in quarrying the stone for the Plymouth breakwater, should
examine the contents of any caverns that he might happen to meet
with. It preceded Dr. Buckland’s exploration of Kirkdale by about four
years.
 In the summer of 1821 a cave was discovered, in a limestone
quarry at Kirkdale, in Yorkshire, which was found to contain bones
and teeth of animals. On hearing of the discovery, Dr. Buckland
posted at once from South Wales to the spot, and published the
result of the explorations in the Philosophical Transactions for the
next year. He brought forward evidence that the cave had been
inhabited by hyænas, and that the broken and gnawed bones of the
rhinoceros, mammoth, stag, bison, and horse belonged to animals
which had been dragged in for food. He also established the fact
that all these animals had lived in Yorkshire in ancient times, and
that it was impossible for the carcases of the hyæna, rhinoceros, and
mammoth to have been floated from those regions where they are
now living into the position where he found their bones. He
subsequently followed up the subject by investigating bone-caves in
Derbyshire, South Wales, and Somerset, as well as in Germany, and
published his great work, “Reliquiæ Diluvianæ,” in 1822, which laid
the foundations of the new science of cave-hunting in this country.
The exploration of Kirkdale followed closely upon that of Gailenreuth,
and was merely the application of those principles of research which
had been discovered in Germany to caves in a new district.
From this time forward bone-caves were discovered in Great
Britain in increasing numbers, and explored by many independent
observers. The famous cavern of Kent’s Hole, near Torquay,
furnished the Rev. J. McEnery, between 1825 and the year 1841, in
which he died, with the first flint implements ever discovered in a
cave along with the bones of extinct animals. He recognized the fact
that they may be proof of the existence of man during the time that
those animals were alive; but the scientific world was not then
sufficiently educated to accept the antiquity of the human race on
the evidence brought forward, and Dr. Buckland himself was so
influenced by the opinions of his times, that he refused even to
entertain the idea. Although the discovery was verified by the
independent researches of Mr. Godwin Austin in 1840, and by the
Torquay Natural History Society in 1846, the force of prejudice was
so strong, that the matter was not thought even worthy of
investigation. Mr. McEnery’s manuscripts were lost until the year
1859, when an abstract of them was published by Mr. Vivian, and
subsequently they were printed in full by Mr. Pengelly, the able
superintendent of the exploration which has been carried on by a
committee of the British Association since 1865, by whom several
thousand flint implements have been obtained, under the conditions
10
pointed out by the Rev. J. McEnery and Mr. Godwin Austen.
While the important question of the antiquity of man was being
passed by as of no account, other caves were being examined in this
country. Those of Banwell, Burrington, Sandford Hill, Bleadon, and
Hutton, in the mountain limestone of the Mendip hills, were being
worked by the Rev. J. Williams and Mr. Beard, and furnished the
magnificent collection of mammalian bones now in the museum at
Taunton. In North Wales, also, Mr. Lloyd discovered a similar suite of
bones in the limestone caves in the neighbourhood of St. Asaph at
Cefn, and in South Wales numerous remains were obtained by many
explorers in those of Pembrokeshire and Gower.
The result of these discoveries was the proof that certain extinct
animals, such as the woolly rhinoceros and the mammoth, had lived
in this country in ancient times, along with two other groups of
species which are at present known only to live in hot and cold
climates—the spotted hyæna and hippopotamus of Africa, with the
reindeer and the marmot of the colder regions of the earth.
The discovery in 1858, and the exploration, of the now famous
cave of Brixham, by the Royal and Geological Societies, marked the
dawn of a new era in cave-hunting. Under the careful supervision of
Mr. Pengelly, flint implements were discovered underneath
stalagmite, and in association with the remains of the hyæna and
woolly rhinoceros and mammoth, in undisturbed red loam, under
conditions that prove man to have been living in Devonshire at the
same time as those animals. This singularly opportune discovery
destroyed for ever the doubts that had overhung the question of the
antiquity of man, and of his co-existence in Europe in company with
the animals whose remains occur both in the caverns and river-
deposits.
In 1847 M. Boucher de Perthes described certain rude flint
implements that he obtained from the fluviatile gravels of Abbeville
(“Antiquités Celtiques,” vol. i.), along with the bones of extinct
animals; and his discovery was treated with the same scepticism in
France as that of the Rev. J. McEnery in England, although it was
verified by flint implements being discovered, under exactly the
same conditions, in the gravels of Amiens, some forty miles away, by
11
Dr. Rigollot. In the autumn of 1858, Dr. Falconer, who had been
superintending the work in the Brixham cave, visited the collection
made by M. de Perthes, while on his way to examine the caves of
Sicily, and recognizing man’s handiwork in the implements, he asked
his friend Mr. Prestwich to explore the Valley of the Somme. This he
accordingly did, and in company with Mr. John Evans, F.R.S., dug out
12
with his own hands an implement from the undisturbed strata, and
thus finally settled the disputed question. It is undoubtedly true, that
scientific opinion was tending towards the acceptance of the
evidence in favour of man having lived in Europe in the Pleistocene
age; but the researches in Brixham cave established the fact on the
highest possible authority, and confirmed the long-neglected
discoveries in the valley of the Somme. By the end of 1859 it was
fully accepted by the scientific world, and caused the exploration of
caves to be carried on with increased vigour.
13
In December 1859, I began the exploration of the hyæna-den
of Wookey Hole, near Wells, Somerset, in company with the Rev. J.
Williamson, and obtained flint instruments along with the remains of
the mammoth, hyæna, woolly rhinoceros, and other animals, under
conditions that proved the contemporaneity of man with the extinct
mammalia. And from that time down to the present date I have
carried on researches in caves in various parts of Great Britain. In
the district of Gower also, many ossiferous caverns were
investigated, in 1858–9–60–1 by Colonel Wood and Dr. Falconer, and
in one of them flint implements were obtained along with the bones
14
of the extinct mammalia. Kent’s Hole, begun in 1865 by the British
Association, and still being worked, furnishes annually a vast number
of bones and teeth of hyænas, rhinoceroses, cave-bears, and
15
horses, and other animals, along with flint and bone implements.
In 1869 I had the good fortune to discover, and subsequently to
explore, a group of sepulchral caves in Denbighshire, which had
been used by an Iberian or Basque race in the Neolithic age
(Chapter V.); and in the following year the Settle Cave Committee
began their work in Yorkshire under my advice. And this has led to
the important conclusion, that a group of caves, extending over a
wide area in the centre and north of England, was occupied by the
Brit-Welsh in the obscure interval which elapsed between the
departure of the Roman legions and the English conquest.
France.—The researches of Buckland into the caves of Great
Britain, and of Goldfuss and others into those of Germany, and more
especially the publication of the “Ossemens Fossiles,” by Cuvier, gave
an impetus to cave-exploration in France which yielded the same
results as in our own country. The mammalia obtained from the cave
of Fouvent (Haut Saone) in 1800 were described in the “Ossemens,”
as well as those from Gondenans. In the Gironde, the Cave of Avison
was explored by M. Billaudel in 1826–27. In the south, Marcel de
Serres, aided by MM. Dubrueil and Jeanjean, examined the
important Cave of Lunel-viel in 1824, and published their results in a
work that holds the same position in France as the “Reliquiæ
Diluvianæ” in England. The caverns of Pondres, Souvignargues, and
of Bize were explored, the two first by M. Christol in 1829, the last
by M. Tournal in 1833, and those of Villefranche (Pyrénées-orient),
Mialet (Gard), and Nabrigas (Lozère) were described by De Serres in
1839, who subsequently added those of Carcas-sonne to the list in
1842. In this year MM. Prevost and J. Desnoyers explored the caves
of Montmorency in the neighbourhood of Paris, and described the
remains discovered in those of Bicêtre. The Cave of Pontil (Hérault)
described by M. de Serres in 1847, was proved in 1864, by Professor
Gervais, to contain two distinct strata, the neolithic lying over the
16
palæolithic, as in Kent’s Hole.
17
In 1860, the famous Cave of Aurignac was proved, by the
investigations of Professor Lartet, to have been inhabited by man in
the life-time of the extinct mammalia. Three years later the caves of
Périgord were explored by that gentleman, along with Mr. Christy,
and yielded results which mark a new era in the history of man in
the remote past. From the remarkable collection of implements and
weapons, the habits and mode of life of the occupants can be
ascertained with tolerable certainty, and from their comparison with
the like articles now in use among savage tribes, it may be
reasonably inferred that they were closely related in blood to the
Eskimos. This most important question will be investigated in its
proper place, in the chapter relating to the palæolithic caves of
France. Professor Lartet, M. Louis Lartet, Sir Charles Lyell, and other
eminent observers believe further, that the interments that have
18
been discovered in Aurignac and in Cro Magnon, in Périgord, are to
be assigned to the same relative age as the occupation of the caves
by man. From the fact, however, that the skeletons in both these
cases were above the strata accumulated by the palæolithic cave-
dwellers, it may be concluded that they were deposited after those
strata were formed, in other words, that they are of a later age.
From 1863 down to the present time very many caves have been
explored in France without any further addition to our knowledge,
excepting the verification of the facts, afforded by the caves of
Brixham and of Périgord, as to the co-existence of man with the
extinct mammalia, and his probable identity in race with the
Eskimos.
19
Belgium.—The caves of Belgium have afforded evidence of
precisely the same nature as those of England and France. Dr.
Schmerling, of Liège, published the results of his researches, begun
in 1829, into the bone-caves on the banks of the Meuse and its
tributaries, in 1833–4, and proved that the mammoth, rhinoceros,
cave-bear, and hyæna formerly lived in that district. He also arrived
at the conclusion that man was living at that remote time, from the
discovery of flint-flakes and human bones along with the remains of
20
those animals in the caves of Engis and Engihoul. In 1853,
Professor Spring discovered a quantity of burned, broken, and cut
bones belonging to women and children, in the Cave of Chauvaux,
which he considered to imply that it had been inhabited by a family
of cannibals. Axes of polished stone were also met with, that
indicated the relative age to be neolithic.
To pass over the human skeleton found in the Neanderthal Cave
in 1857 by Dr. Fuhlroth, which is of doubtful antiquity, the next
discoveries of importance are those made by M. Dupont in the years
1864–70, in the province of Namur, that established the fact that the
same race of men who inhabited Auvergne in the palæolithic age
had also lived in Belgium. M. Dupont considers that the interments
21
in the Trou de Frontal belong also to the palæolithic age, and that
therefore man at that remote time was possessed of religious ideas.
Before, however, this view can be accepted, it will be necessary to
show the exact relation of the bones of the reindeer, chamois,
mammoth, and other animals found outside the slab of stone, at the
mouth of the sepulchral chamber, to the human remains within. In
this case, as in Aurignac and Cro Magnon, the evidence seems to me
insufficient to establish so important a conclusion.
Southern Europe.—In southern Europe the bone-caves of Sicily,
worked in 1829 for the sake of the animal remains to be used in
sugar refining, were scientifically examined by Dr. Falconer in 1859;
those of Malta by Captain Spratt in the same year; and those of
Gibraltar by Captain Broome in the years 1862–8. They established
the existence of the serval and the African elephant, and other
characteristic African species, in Europe, and offer as we shall see in
this work, important testimony as to the geography of the
Mediterranean area in the Pleistocene age.
 In this outline of the history of cave-exploration it will be seen,
that the additions to our knowledge of the past have been neither
few nor insignificant, nor in one line of inquiry. And if the attention
which is now being directed to the subject be due to the general
development of scientific thought, it is equally true, that the results
have reacted on scientific thought in general, and have especially
benefited the sciences of geology, archæology, and history. A rich
field of investigation lies before the cave-hunter, in Greece, Palestine,
Lycia, Persia, and the limestone plateaux of central Asia; and since
these discoveries have been so valuable in central and north-western
Europe, what may we not recover from the grasp of oblivion, of the
infancy and early culture of mankind in the very birth-place and
“pathway of the nations”?
 CHAPTER II.
PHYSICAL HISTORY OF CAVES.

Caves formed by the Sea and by Volcanic Action.—Caves in Arenaceous

Rocks.—Caves in Calcareous Rocks of various ages.—Their Relation to
Pot-holes, “Cirques,” and Ravines.—The Water-cave of Wookey Hole.—
The Goatchurch Cave.—The Water-caves of Derbyshire.—Of Yorkshire.
—The Ingleborough Cave.—The Rate of Deposit of Stalagmite.—The
Descent into Helln Pot.—The Caves and Pots round Weathercote.—The
Formation of Caves, Pot-holes, and Ravines.—Caverns not generally
formed in line of Faults.—Of various Ages.—Their Filling-up.—The Cave
of Caldy.—The Blackrock Cave.—Great quantity of Carbonate of Lime
dissolved by Atmospheric Water.—The Circulation of Carbonate of
Lime.—The Temperature of Caves.—Conclusion.

Caves formed by the Sea and by Volcanic Action.

In this chapter we shall treat of the origin of caves and of their place
in physical geography. The most obvious agent in hollowing out
caves is the sea. The set of the current, the tremendous force of the
breakers, and the grinding of the shingle, inevitably discover the
weak places in the cliff, and leave caves as the results of their work,
modified in each case by the local conditions of the rock. Caves
formed in this manner have certain characters which are easily
recognized. Their floors are very rarely much out of the horizontal,
their outlook is over the sea, and they very seldom penetrate far into
the cliff. A general parallelism is also to be observed in a group in
the same district, and their entrances are all in the same horizontal
plane, or in a succession of horizontal and parallel planes. In some
cases they are elevated above the present reach of the waves, and
mark the line at which the sea formerly stood. From their generally
inaccessible position sea-caves have very rarely been occupied by
man, and the history of their formation is so obvious that it requires
no further notice. Among them the famous Fingal’s Cave, off the
north coast of Ireland, and that of Staffa, on the opposite shore of
Scotland, hollowed out of columnar basalt, are perhaps the most
remarkable in Europe.
In volcanic regions also there are caves formed by the passage
of lava to the surface of the ground, or by the imprisoned steam and
gases in the lava while it was in a molten state: but these are of
comparatively little importance so far as relates to the general
question of caves, from the very small areas which are occupied by
active volcanoes in Europe. They have been observed in Vesuvius,
Etna, Iceland, and Teneriffe.

Caves in Arenaceous Rocks.

Caves also occur sometimes in sandstones, in which case they
are the result of the erosion of the lines of the joints by the passage
of subaërial water, and if the joints happen to traverse a stratum less
compacted than the rest, the weak point is discovered, and a hollow
is formed extending laterally from the original fissure. The massive
millstone grit of Derbyshire and Yorkshire present many examples of
this, as for instance in Kinderscout in the former county. The rocks at
Tunbridge Wells also show to what extent the joints in the Wealden
sandstones may become open fissures, more or less connected with
caves, on a small scale, by the mere mechanical action of water. M.
Desnoyers gives instances of the same kind in the Tertiary
sandstones of the Paris basin, which have furnished remains of
rhinoceros, reindeer, hyæna, and bear. Caverns, however, in the
sandstone are rarely of great extent, and may be passed over as
being of small importance in comparison with those in the
calcareous rocks.

Caves in Calcareous Rocks of various ages.

It has long been known that wherever the calcareous strata are
sufficiently hard and compact to support a roof, caves are to be
found in greater or less abundance. Those of Devonshire occur in
the Devonian limestone; those of Somerset, Nottinghamshire,
Yorkshire, Derbyshire, and Northumberland, as well as of Belgium
and Westphalia, in that of the carboniferous age. In France also,
those of Maine and Anjou, and most of those of the Pyrenees and in
the department of Aude, are hollowed in carboniferous limestone, as
well as the greater part of those in North America, in Virginia, and
Kentucky. The cave of Kirkdale in Yorkshire, and most of those in
Franconia and in Bavaria penetrate Jurassic limestones, which have
received the name of Hohlenkalkstein from the abundance of
caverns which they contain. They are developed on a large scale in
the Swiss and French Jura, and in some cases afford passage to
powerful streams, and in others are more or less filled with ice, thus
constituting the singular “glacières” that have been so ably explored
22
by the Rev. G. F. Browne.
The compact Neocomian and Cretaceous limestones contain
most of the caverns of Périgord, Quercy, and Angoumois, and some
of those in Provence and Languedoc, those of Northern Italy, Sicily,
Greece, Dalmatia, Carniola, and Turkey in Europe, of Asia Minor and
Palestine.
23
The tertiary limestones, writes M. Desnoyers, offer sometimes,
but very rarely, caves that have become celebrated for the bones
which they contain, such as those of Lunel-Viel, near Montpelier,
those of Pondres and Souvignargues, near Sommières (Gard), and of
Saint Macaire (Gironde). The same may also be said of the calcaire
grossier of the basin of Paris.
Certain rocks composed of gypsum also contain caverns of the
same sort as those in the limestones. In Thuringia, for example,
near Eisleben, they occur in the saliferous and gypseous strata of
the zechstein, and are connected with large gulfs and cirques on the
surface, which are sometimes filled with water. In the
neighbourhood of Paris, and especially at Montmorency, they contain
numerous bones of the extinct mammalia. M. Desnoyers points out
their identity, in all essentials, with those in calcareous strata, and
infers that they have been produced in the same way. Some of them
may have been formed by the removal of the salt, which is very
frequently interbedded with the gypsum, by the passage of water. In
Cheshire the pumping of the brine from the saliferous and gypseous
strata produces subterranean hollows, which sometimes fall in and
eventually cause depressions on the surface, such as those which
are now destroying the town of Northwich, and causing the
neighbouring tidal estuary to extend over what was formerly
meadow land. This explanation, however, will not apply to those in
the neighbourhood of Paris, because there is no trace of their ever
having contained salt.

The Relation of Caves to Pot-holes, “Cirques,” and

Ravines.
The caverns hollowed in calcareous rocks present features by
which they are distinguished from any others. They open, for the
most part, on the abrupt sides of valleys and ravines at various
levels, being arranged round the main axis of erosion just as
branches are arranged round the trunk of a tree—as, for example, in
Cheddar Pass. The transition in some cases from the valley to the
ravine, and from the ravine to the cave, is so gradual, that it is
impossible to deny that all three are due to the same cause. The
caves themselves ramify in the same irregular fashion as the valleys,
and are to be viewed merely as the capillaries in the general valley
system, through which the rainfall passes to join the main channels.
Very frequently, however, the drainage has found an outlet at a
lower level, and its ancient passage is left dry; but in all cases
unmistakeable proof of the erosive action of water is to be seen in
the sand, gravel, and clay which compose the floor, as well as in the
worn surfaces of the sides and the bottom.
In all districts in which caves occur are funnel-shaped cavities of
various sizes, known as “pot-holes” or “swallow-holes” in Britain, as
“betoires,” “chaldrons du diable,” “marmites de géants,” in France,
and as “kata-vothra” in Greece, in which the rainfall is collected
before it finally disappears in the subterranean passages. They are
to be seen in all stages; sometimes being mere shallow funnels, that
only contain water after excessive rain, and at others as profound
vertical shafts, into which the water is continually falling, as in Helln
Pot, in Yorkshire. The cirques, also, described by M. Desnoyers,
belong to the same class of cavities, although all those which are
24
mentioned by the Rev. T. G. Bonney, at the head of valleys, and in
some cases hollowed in shale and igneous rocks, are most probably
to be referred to the vertical, chisel-like action of streams flowing
under physical conditions, that resemble those under which the
cañons of the Colorado, or of the Zambesi, are being excavated, and
in which frost, ice, and snow have played a very subordinate part.
The intimate relation between pot-holes, caves, ravines, and
valleys will be discussed in the rest of this chapter, and illustrated by
English examples; and then we shall proceed to show that the
chemical action of the carbonic acid in the rain-water, and the
mechanical friction of the sand and gravel, set in motion by the
water, by which Professor Phillips explains the origin of caves, will
equally explain the pot-holes and ravines by which they are
invariably accompanied.
 The Water-Cave of Wookey Hole, near Wells,
Somerset.
Caves may be divided into two classes: those which are now
mere passages for water, in which the history of their formation may
be studied, and those which are dry, and capable of affording shelter
to man and the lower animals. Among the water-caves, that of
25
Wookey Hole is to be noticed first, since its very name implies that
it was known to the Celtic inhabitants of the south of England, and
since it was among the first, if not the first, of those examined with
26
any care in this country, Mr. John Beaumont having brought it
before the notice of the Royal Society in the year 1680.
The hamlet of Wookey Hole nestles in a valley, through which
flows the river Axe, and the valley passes insensibly, at its upper
end, into a ravine, which is closed abruptly by a wall of rock (Fig. 1),
about two hundred feet high, covered with long streamers and
festoons of ivy, and affording scanty hold, on its ledges and in its
fissures, to ferns, brambles, and ash saplings. At its base the river
Axe issues, in full current, out of the cave, the lower entrance of
which it completely blocks up, since the water has been kept back by
a weir, for the use of a paper-mill a little distance away. A narrow
path through the wood, on the north side of the ravine, leads to the
27
only entrance now open. Thence a narrow passage leads
downward into the rock, until, suddenly, you find yourself in a large
chamber, at the water level. Then you pass over a ridge, covered
with a delicate fretwork of dripstone, with each tiny hollow full of
water, and ornamented with brilliant lime crystals. One shapeless
mass of dripstone is known in local tradition as the Witch of Wookey,
turned into stone by the prayers of a Glastonbury monk. Beyond this
the chamber expands considerably, being some seventy or eighty
feet high, and adorned with beautiful stalactites, far out of the reach
of visitors. The water, which bars further entrance, forms a deep
pool, which Mr. James Parker managed to cross on a raft (see
Appendix I.) into another chamber, which was apparently easy of
access before the construction of the weir. It was in this further
chamber that Dr. Buckland found human remains and pottery.

Fig. 1.—Diagram of Wookey Hole Cave and Ravine.

The cave has been proved to extend as far as the village of

Priddy, about two miles off, on the Mendip hills, by the fact observed
by Mr. Beaumont, that the water used in washing the lead ore at
that spot, in his time, found its way into the river Axe, and poisoned
cattle in the valley of Wookey. And this observation has been verified
during the last few years by throwing in colour and chopped straw.
The stream at Priddy sinks into a swallow-hole (Fig. 1), and has its
subterranean course determined by the southerly dip of the rock, by
which the joints running north and south afford a more free passage
to the water than those running east and west. The cave is merely a
subterranean extension of the ravine in the same line, as far as the
swallow-hole, and all three have been hollowed, as we shall see
presently, by the action of the stream and of carbonic acid in the
water.

The Goatchurch Cave.

 The largest cavern in the Mendip hills is that locally known as
the Goatchurch, which opens on the eastern side of the lower of the
two ravines that branch from the magnificent defile of Burrington
Combe, about two miles from the village of Wrington, at the height
of about 120 feet from the bottom of the ravine. After creeping
along a narrow, muddy passage, with a steep descent to the west,
at an angle of about 30°, you suddenly pass into a stalactitic
chamber of considerable height and size. From it two small vertical
shafts lead into the lower set of chambers and passages; the first
being blocked up, and the second being close to a large barrel-
shaped stalagmite, to which Mr. Ayshford Sanford, Mr. James Parker,
and myself fastened our ropes when we explored the cave in 1864.
The latter affords access into a passage, beautifully arched, and
passing horizontally east and west, and just large enough to admit a
man walking upright. At the further end numerous open fissures,
caused by the erosion of the joints in the limestone, cross it at right
angles, and pass into several ill-defined chambers, partially
stalactitic, but for the most part filled with loose, bare, cubical
masses of limestone. Two of the transverse fissures lead into a large
chamber, at a lower level. At its lower end, on crawling along a
narrow passage, we came into a second chamber, also of
considerable height and depth, at the bottom of which the noise of
flowing water can be heard through two vertical holes, just large
enough to admit of access. On sliding down one of these we found
ourselves in a third chamber, which was traversed by a subterranean
stream, doubtless in part the same which disappears in the ravine,
at a point eighty feet above by aneroid measurement. The
temperature of the water, as compared with that of the stream
outside (49° : 59°), renders it very probable that, between the point
of disappearance in the ravine and reappearance in the cave, it is
joined by a stream of considerable subterranean length, since the
water could not have lost ten degrees in the short interval which it
had to traverse, were it supplied only from the stream in the ravine.
From the point of its disappearance in the cave, the water passes
downwards to join the main current flowing underneath Burrington
Combe, that gushes forth in great volume at Rickford. The lowest
portion of the cave was eighteen or twenty feet below the stream,
and 220 feet below the entrance of the cavern.
On examining the floors of the chambers and passages, we
discovered that they were composed of the same kind of sediment
as that which is now being deposited by the water in Wookey Hole,
and there could be no doubt but that they had been originally
traversed by water. For this to have taken place it is necessary to
suppose that, while the Goatchurch was a water cave, the ravine on
which it opens was not deeper than the entrance—in other words,
that in the interval between the formation and excavation of the
chambers and passages, to the present time, the ravine has been
excavated in the limestone to a depth of a hundred and twenty feet,
and the water which originally passed through the entrance has
found its way, by a new series of passages, to the point where it
appears at the bottom of the cave.
We obtained evidence that the horizontal passage, immediately
below the first vertical descent, had been inhabited at a very remote
period. At the spot where Mr. Beard, of Banwell, obtained a fine tusk
of mammoth, we found a molar of bear, and a fragment of flint,
which were imbedded in red earth, and were underneath a crust of
stalagmite of about two inches in thickness. It would follow from
this, that the date of the formation of this part of the cave was
before the time when the traces of elephants, bears, and of man
were introduced.
The cave is the resort of numerous badgers. On hiding ourselves
in one of the transverse fissures, and throwing our light across the
horizontal passage, these animals ran to and fro across the lighted
field with extraordinary swiftness, and had it not been for the white
streaks on the sides of their heads, which flashed back the light,
they would not have been observed. Though they are rarely caught,
they must be abundant in the district.
Like all the other large caverns in the district, it has its legends.
The dwellers in the neighbourhood, who have never cared to explore
its recesses, relate that a certain dog put in here found its way out,
after many days, at Wookey Hole, having lost all its hair in
scrambling through the narrow passages. At Cheddar the same
legend is appropriated to the Cheddar cave. At Wookey the dog is
said to have travelled back to Cheddar. Some eighteen years ago,
while exploring the limestone caves at Llanamynech, on the English
border of Montgomeryshire, I met with a similar story. A man playing
the bagpipes is said to have entered one of the caves, well
provisioned with Welsh mutton, and after he had been in for some
time his bagpipes were heard two miles from the entrance,
underneath the small town of Llanamynech. He never returned to
tell his tale. The few bones found in the cave are supposed to be
those which he had picked on the way. This is doubtless another
form of the story of the dog; both owe their origin to the vague
impression, which most people have, of the great extent of caverns,
and both versions are equally current in France and Germany.

The Water-caves of Derbyshire.

The celebrated cavern of the Peak, at Castleton in Derbyshire,
presents the same essential character as that of Wookey Hole. It
runs into the hill-side at the end of the ravine, and is traversed by a
powerful stream of water, which has been met with in driving an
horizontal adit in lead-mining at a considerable distance from the
entrance, and finally traced to a distant swallow-hole. At a little
distance from Buxton a smaller cave, known as Poole’s Cavern, is in
part traversed by water, which has found an outlet at a lower level,
and allowed of the present entrance being used by the Brit-Welsh
(Romano-Celtic) inhabitants of the district as a habitation in the fifth
28
and sixth centuries. There are, besides these, very many others,
some known, others unknown, that debouch on the sides of the
dales in Derbyshire and Staffordshire, and are all well worthy of
examination, since they illustrate not merely the history of the
formation of caves, but also have been proved to contain works of