Elements of Ecology PDF
Preview Elements of Ecology
ELEMENTS OF E C O LO GY CANADIAN EDITION THOMAS M. SMITH ROBERT LEO SMITH ISOBEL WATERS Toronto Vice-President, Editorial Director: Gary Bennett Proofreader: Dawn Hunter Senior Acquisitions Editor: Lisa Rahn Compositor: Aptara®, Inc. Senior Marketing Manager: Kim Ukrainec Photo and Permissions Researcher: Monika Senior Developmental Editor: John Polanszky Schurmann Project Manager: Jessica Hellen Art Director: Julia Hall Manufacturing Specialist: Jane Schell Cover and Interior Designer: Miriam Blier Production Editor: Niraj Bhatt, Aptara®, Inc. Cover Image: Jason Carter / Wapati at Ha Ling Copy Editor: Julia Cochrane Credits and acknowledgments of material borrowed from other sources and reproduced, with permission, in this textbook appear on the appropriate page within the text and on p. C-1. Original edition published by Pearson Education, Inc., Upper Saddle River, New Jersey, USA. Copyright © 2012 Pearson Education, Inc. This edition is authorized for sale only in Canada. If you purchased this book outside the United States or Canada, you should be aware that it has been imported without the approval of the publisher or author. Copyright © 2014 Pearson Canada Inc. All rights reserved. Manufactured in the United States of America. This publication is protected by copyright and permission should be obtained from the publisher prior to any prohibited reproduction, storage in a retrieval system, or transmission in any form or by any means, electronic, mechanical, photocopying, recording, or likewise. To obtain permission(s) to use material from this work, please submit a written request to Pearson Canada Inc., Permissions Department, 26 Prince Andrew Place, Don Mills, Ontario, M3C 2T8, or fax your request to 416-447-3126, or submit a request to Permissions Requests at w ww.pearsoncanada.ca . 10 9 8 7 6 5 4 3 2 1 [CKV] Library and Archives Canada Cataloguing in Publication Smith, T. M. (Thomas Michael), 1955- Elements of ecology / Thomas M. Smith, Robert Leo Smith and Isobel Waters. - 1st ed. Includes bibliographical references and index. ISBN 978-0-321-51201-7 1. Ecology. I. Smith, Robert Leo II. Waters, Isobel III. Title. QH541.S65 2012 577 C2012-903757-5 Environmental Statement This book is carefully crafted to minimize environmental impact. Pearson Canada is proud to report that the materials used to manufacture this book originated from sources committed to sustainable forestry practices, tree harvesting, and associated land management. The binding, cover, and paper come from facilities that minimize waste, energy usage, and the use of harmful chemicals. Equally important, Pearson Canada closes the loop by recycling every out-of-date text returned to our warehouse. We pulp the books, and the pulp is used to produce other items such as paper coffee cups or shopping bags. The future holds great promise for reducing our impact on Earth’s environment, and Pearson Canada is proud to be leading the way in this initiative. From production of the book to putting a copy in your hands, we strive to publish the best books with the most up-to-date and accurate content, and to do so in ways that minimize our impact on Earth. ISBN 978-0-321-51201-7 Brief Contents Preface xx Chapter 1 The Nature of Ecology 1 PART 1 The Physical Environment 18 Chapter 2 Climate 20 Chapter 3 The Aquatic Environment 39 Chapter 4 The Terrestrial Environment 57 PART 2 The Organism and Its Environment 74 Chapter 5 Ecological Genetics 76 Chapter 6 Plant Adaptations to the Environment 100 Chapter 7 Animal Adaptations to the Environment 126 Chapter 8 Life History Patterns 158 PART 3 Populations 180 Chapter 9 Properties of Populations 182 Chapter 10 Population Growth 201 Chapter 11 I ntraspecific Population Regulation 221 Chapter 12 M etapopulations 239 PART 4 Species Interactions 254 Chapter 13 I nterspecific Competition 256 Chapter 14 P redation 279 Chapter 15 P arasitism and Mutualism 308 PART 5 Community Ecology 330 Chapter 16 C ommunity Structure 332 Chapter 17 C ommunity Dynamics 362 Chapter 18 L andscape Ecology 385 PART 6 Ecosystem Ecology 4 06 Chapter 19 S YSTEMS ECOLOGY 408 Chapter 20 E cosystem Energetics 427 Chapter 21 D ecomposition and Nutrient Cycling 447 Chapter 22 B iogeochemical Cycles 473 iii iv BRIEF CONTENTS PART 7 E cological Biogeography 496 Chapter 23 T errestrial Ecosystems 498 Chapter 24 A quatic Ecosystems 527 Chapter 25 Coastal and Wetland Ecosystems 547 PART 8 H uman Ecology 564 Chapter 26 P opulation Growth and Sustainability 566 Chapter 27 C onservation Ecology 593 Chapter 28 G lobal Climate Change 622 References R-1 Glossary G-1 Credits C-1 Index I-1 Contents Preface xx Chapter 1 The Nature of Ecology 1 1.1 ECOLOGY DEFINED: Ecology Studies Organisms Interacting with Their Environment 2 1.2 HISTORY OF ECOLOGY: Ecology Is Rooted in Many Scientific Disciplines 3 1.3 ORGANISMS AND ECOSYSTEMS: Organisms Interact with the Environment within Ecosystems 3 1.4 ECOLOGICAL HIERARCHY: Ecology Operates at Many Levels, from the Individual to the Biosphere 4 1.5 IMPORTANCE OF SCALE: Ecologists Study Pattern and Process at Different Scales 5 1.6 ECOLOGY AS A SCIENCE: Ecologists Investigate Nature by Using the Scientific Method 6 (cid:2) QUANTIFYING ECOLOGY 1.1: Classifying Ecological Data 8 1.7 MODELS IN ECOLOGY: Mathematical and Descriptive Models Allow Predictions 9 1.8 UNCERTAINTY AND VARIATION: Ecological Phenomena Exhibit Uncertainty and Variation 10 (cid:2) QUANTIFYING ECOLOGY 1.2: Displaying Ecological Data 11 (cid:2) ECOLOGICAL ISSUES: The Human Factor 13 1.9 AN INTERDISCIPLINARY SCIENCE: Ecology Has Strong Ties to Many Other Fields 14 1.10 DUAL FOCUS: Both the Individual and the Ecosystem Are Basic Units of Nature 14 Summary 15 • Key Terms 15 • Study Questions 16 • Further Readings 16 PART 1 The Physical Environment 18 Chapter 2 Climate 20 2.1 SOLAR RADIATION: Solar Energy Is Subject to Various Fates Before It Is Intercepted by Earth 21 2.2 RADIATION AND TEMPERATURE: Latitudinal and Seasonal Radiation Patterns Affect Temperature 23 2.3 ELEVATION AND TEMPERATURE: Air Temperature in the Troposphere Decreases with Elevation 24 2.4 ATMOSPHERIC CIRCULATION: Global Circulation of Air Masses Affects Climate 26 2.5 OCEAN CURRENTS: Solar Energy, Wind, and Earth’s Rotation Create Ocean Currents 27 2.6 HUMIDITY: Temperature Influences the Moisture Content of Air 28 2.7 PRECIPITATION: Movements of Air Masses Create Global Precipitation Patterns 29 2.8 SNOW AND ICE: Frozen Precipitation Has Many Ecological Effects 31 2.9 EFFECTS OF TOPOGRAPHY: Landforms Influence Regional and Local Precipitation 31 2.10 IRREGULAR CLIMATIC VARIATION: Earth Experiences Significant Irregular Variation in Climate 31 v vi CONTENTS 2.11 MICROCLIMATES: Local Microclimates Define the Conditions Experienced by Individuals 33 (cid:2) ECOLOGICAL ISSUES: Urban Microclimates 34 Summary 36 • Key Terms 37 • Study Questions 37 • Further Readings 37 Chapter 3 The Aquatic Environment 39 3.1 PROPERTIES OF WATER: The Unique Properties of Water Determine Its Ecological Importance 40 3.2 LIGHT IN WATER: Light Quantity and Quality Vary with Water Depth 42 3.3 WATER TEMPERATURE: The Effect of Water Depth on Temperature Varies Seasonally 42 3.4 WATER AS A SOLVENT: Many Substances Dissolve Readily in Water 4 4 3.5 OXYGEN IN WATER: Abiotic and Biotic Factors Affect the O Content 2 of Water 44 (cid:2) RESEARCH IN ECOLOGY: The Extreme Environment of Hydrothermal Vents 46 3.6 WATER ACIDITY: A Complex Sequence of Carbon Reactions Affects Aquatic pH 48 3.7 WATER CYCLE: Water Circulates between Earth and the Atmosphere 48 3.8 WATER MOVEMENTS: Currents and Waves Shape Freshwater and Marine Habitats 50 (cid:2) ECOLOGICAL ISSUES: Our Disappearing Groundwater Resources 51 3.9 TIDES: Tide Action Dominates Marine Coastal Habitats 53 3.10 ESTUARIES: Transition Areas between Freshwater and Saltwater Are Unique Habitats 53 Summary 54 • Key Terms 55 • Study Questions 55 • Further Readings 56 Chapter 4 The Terrestrial Environment 57 4.1 LIFE ON LAND: Terrestrial Habitats Present Challenges and Opportunities 58 4.2 LIGHT AND VEGETATION: Vegetative Cover Influences the Vertical Light Profile 59 (cid:2) QUANTIFYING ECOLOGY 4.1: Measuring Light Attenuation 61 4.3 SOIL DEFINED: Soil Is the Foundation upon Which All Terrestrial Life Depends 62 4.4 SOIL FORMATION: Weathering Initiates Soil Formation, Which Is Affected by Six Factors 63 4.5 SOIL PROPERTIES: Soils Differ in Physical Traits 64 4.6 SOIL HORIZONS: Soil Has Distinct Horizontal Layers 64 4.7 SOIL MOISTURE: Water-Holding Capacity Is a Vital Property of Soils 66 4.8 SOIL FERTILITY: Ion Exchange Capacity Affects Soil Fertility 66 (cid:2) ECOLOGICAL ISSUES: Have We Passed Peak Soil? 67 4.9 SOIL TYPES: Different Processes Produce Different Soil Types 69 Summary 71 • Key Terms 72 • Study Questions 72 • Further Readings 72 PART 2 T he Organism and Its Environment 74 Chapter 5 Ecological Genetics 76 5.1 ADAPTATIONS: Natural Selection Promotes Adaptive Traits 77 5.2 GENES: Adaptations Are Coded by Genes, the Basic Units of Inheritance 77 5.3 PHENOTYPES: Genotype Expression Is Affected by the Environment 78 CONTENTS vii 5.4 GENETIC VARIATION: Allele and Genotype Frequencies Differ within and among Populations 79 5.5 TYPES OF NATURAL SELECTION: Natural Selection Can Be Directional, Stabilizing, or Disruptive 79 5.6 EVOLUTIONARY PROCESSES: Various Processes Disrupt the Hardy– Weinberg Equilibrium 81 (cid:2) ECOLOGICAL ISSUES: Selecting for Antibiotic Resistance 82 5.7 GENETIC DIFFERENTIATION: Natural Selection Alters Genetic Variation among Populations of Widely Distributed Species 84 5.8 TRADE-OFFS AND CONSTRAINTS: Adaptations Entail Ecological Compromises 85 (cid:2) RESEARCH IN ECOLOGY: Dynamic Evolution in Sticklebacks 87 5.9 SPECIATION: Evolution Can Lead to the Emergence of New Species 89 5.10 HYBRIDIZATION: Hybridization Can Prevent or Promote Speciation 91 5.11 ABSENCE OF PERFECTION: No Organism Is Perfectly Adapted to Its Environment 93 5.12 PHENOTYPIC PLASTICITY: Organisms Respond to Environmental Variation at the Individual and Population Levels 94 Summary 97 • Key Terms 98 • Study Questions 98 • Further Readings 99 Chapter 6 Plant Adaptations to the Environment 100 6.1 CARBON METABOLISM: Photosynthesis Converts CO into Simple 2 Sugars 101 6.2 IMPACT OF LIGHT: Photosynthetic Rate Varies with Light Intensity 102 6.3 GAS EXCHANGE: Plants Exchange CO and HO Vapour with the 2 2 Atmosphere 103 6.4 WATER MOVEMENT: Water Follows a Free-Energy Gradient from the Soil through the Plant to the Atmosphere 103 6.5 AQUATIC PHOTOSYNTHESIS: Some Aquatic Photoautotrophs Utilize Bicarbonate 106 6.6 PLANT TEMPERATURE: Plants Exchange Heat Energy with Their Environment 106 6.7 CARBON ALLOCATION: Plants Allocate Net Photosynthetic Carbon G ain to New Tissues 107 6.8 INTERDEPENDENCE OF ADAPTATIONS: Plant Adaptations Reflect Responses to Multiple Factors 108 6.9 ADAPTATIONS TO LIGHT: Shade Tolerance Reflects Genotype and Acclimation 109 (cid:2) QUANTIFYING ECOLOGY 6.1: Calculating Plant Growth Rate 111 6.10 ADAPTATIONS TO DROUGHT: Many Adaptations Help Plants Cope with Limited Water 113 6.11 ADAPTATIONS TO TEMPERATURE: Plants Have Various Adaptations for Surviving Extreme Temperatures 116 6.12 ADAPTATIONS TO MINERALS: Conservative Strategies Help Plants Tolerate Infertile Sites 118 6.13 ADAPTATIONS TO FLOODING: Submerged Substrates Impose O Stress 2 on Plants 121 6.14 PERIODICITY: Many Plants Have Predictable Daily and Seasonal Activity Patterns 122 Summary 123 • Key Terms 124 • Study Questions 124 • Further Readings 125 viii CONTENTS Chapter 7 Animal Adaptations to the Environment 126 7.1 BODY SIZE: Size Imposes a Fundamental Constraint on Evolution 127 7.2 FOOD ACQUISITION: Animals Acquire Energy and Nutrients by Different Strategies 129 7.3 NUTRITIONAL REQUIREMENTS: Animals Meet Most Nutritional Needs from Their Diet 131 7.4 OXYGEN UPTAKE: Structural Adaptations Facilitate O Uptake 133 2 7.5 HOMEOSTASIS: Regulation of Internal Conditions Involves Negative Feedback 134 7.6 THERMAL EXCHANGE: Animals Exchange Heat Energy with Their Environment 134 (cid:2) QUANTIFYING ECOLOGY 7.1: Heat Exchange and Temperature Regulation 136 7.7 POIKILOTHERMY: Poikilotherms Rely on External Heat Sources 138 7.8 HOMEOTHERMY: Homeotherms Reduce Thermal Constraints by Regulating Body Temperature 140 7.9 TRADE-OFFS IN THERMAL REGULATION: Endothermy and Ectothermy Have Benefits and Costs 141 (cid:2) RESEARCH IN ECOLOGY: Prehistoric Snakes as Paleothermometers 143 7.10 HETEROTHERMY: Some Species Practise Both Ectothermy and Endothermy 145 7.11 TEMPERATURE EXTREMES: Animals Have Adaptations for Extreme Heat or Cold 147 7.12 SNOW: Adaptations Allow Animals to Benefit from Snow Cover and Reduce Its Negative Impacts 149 7.13 WATER BALANCE: Terrestrial Animals Are Constrained by Water Uptake and Loss 149 7.14 OSMOTIC BALANCE: Aquatic Animals Face Problems of Water and Salt Balance 150 7.15 WATER DEPTH: Aquatic Animals Have Specialized Adaptations to Deep Water 151 7.16 PERIODICITY: Daily and Seasonal Light Cycles Influence Animal Activity 151 Summary 154 • Key Terms 156 • Study Questions 156 • Further Readings 157 Chapter 8 Life History Patterns 158 8.1 TYPES OF REPRODUCTION: Reproduction May Be Sexual or Asexual 159 8.2 MODES OF SEXUAL REPRODUCTION: Sexual Organisms Are Unisexual or Hermaphroditic 160 8.3 MATING SYSTEMS: Resource Supply Affects the Adaptive Value of Mating Systems 162 8.4 SEXUAL SELECTION: Acquiring Mates Involves Intrasexual and/or Intersexual Selection 163 (cid:2) RESEARCH IN ECOLOGY: Sexual Conflict and the Coevolutionary Arms Race 165 8.5 IMPACT OF RESOURCES: Females May Choose Mates Based on Resource Availability 167 8.6 REPRODUCTIVE EFFORT: Species Vary in the Time and Energy Allocated to Reproduction 168 (cid:2) QUANTIFYING ECOLOGY 8.1: Interpreting Trade-Offs 170 8.7 REPRODUCTIVE TIMING: Differences in Timing of Reproduction Entail Trade-offs 171 8.8 REPRODUCTIVE OUTPUT: Parental Traits and Food Supply Affect Fecundity 171
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