Ecological Methodology Second Edition

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1 Ecological Methodology Second Edition Charles J. Krebs University of British Columbia Technische Universitat Darmstadt FACHBEREICH 10 BIOLOGIE Bi bliothek SchnittspahnstraBe 10 D Darmstadt Inv.-Nr. An imprint of Addison Wesley Longman, Inc. Menlo Park, California Reading, Massachusetts New York Harlow, England Don Mills, Ontario Amsterdam Madrid Sydney Mexico City

2 Contents Chapter 1 Ecological Data 1.1 Designing Field Studies 1.2 Scales of Measurement 1.3 Statistical Inference 1.4 Data Records 12 Questions and Problems Part One Estimating Abundance in Animal and Plant Populations 17 Chapter 2 Estimating Abundance: Mark-Recapture Techniques Petersen Method Confidence Intervals Sample Size Estimation Assumptions of the Petersen Method Schnabel Method Confidence Intervals Assumptions of the Schnabel Method Jolly-Seber Method Confidence Intervals Assumptions of the Jolly-Seber Method ^' Tests of Equal Catchability Zero-Truncated Poisson Test Chapman's Test Leslie's Test Leslie, Chitty, and Chitty Test Planning a Mark-Recapture Study What to Do If Nothing Works Summary 65 Questions and Problems 66 Chapter 3 Estimating Abundance: Removal Methods and Resight Methods Exploited Population Techniques Change-in-Ratio Methods Eberhardt's Removal Method Catch-Effort Methods Resight Methods Computer Programs for Population Estimators Enumeration Methods 93

3 vi Contents Estimating Density Boundary Strip Methods Nested Grids Method Trapping Web Method Summary 100 Questions and Problems Chapter Chapter Estimating Abundance: Quadrat Counts 105 Quadrat Size and Shape Wiegert's Method Hendricks's Method When Should You Ignore These Recommendations? 113 Statistical Distributions Poisson Distribution Negative Binomial Distribution 123 Line Intercept Method 139 Aerial Surveys of Wildlife Populations " Correcting for Bias in Aerial Surveys Sampling in Aerial Surveys 146 Summary 154 Questions and Problems 154 Estimating Abundance: Line Transects and Distance Methods 158 Line Intersects Hay ne Estimator Fourier Series Estimator Shape-Restricted Estimator 167 Distance Methods Byth and Ripley Procedure T-Square Sampling Procedure Ordered Distance Method Variable-Area Transect Method Point-Quarter Method 182 Summary 184 Questions and Problems 185 Part Two Spatial Pattern in Animal and Plant Populations 189 Chapter 6 Spatial Pattern and Indices of Dispersion Methods for Spatial Maps Nearest-Neighbor Methods Distances to Second-nth Nearest Neighbors More Sphisticated Techniques for Spatial Maps Contiguous Quadrats Testing for Spatial Pattern Spatial Pattern from Distance Methods Byth and Riply Procedure T-Square Sampling Procedure Eberhardt's Test Variable-Area Transects 211

4 Contents VII 6.4 Indices of Dispersion for Quadrat Counts Variance-to-Mean Ratio k of the Negative Binomial Green's Coefficient Morisita's Index of Dispersion Standardized Morisita Index Distance-to-Regularity Indices 6.5 Summary 223 Questions and Problems , Part Three Sampling and Experimental Design 227 Chapter 7 Sample Size Determination and Statistical Power Sample Size for Continuous Variables Means from a Normal Distribution Comparison of Two Means Variances from a Normal Distribution Sample Size for Discrete Variables Proportions and Percentages Counts from a Poisson Distribution Counts from a Negative Binomial Distribution Sample Size for~specialized Ecological Variables Mark-Recapture Estimates Line Transect Estimates Distance Methods Change^in-Ratio Methods Statistical Power Analysis Estimates of Effect Size for Continuous Variables Effect Size for Categorical Variables PowerAnalysis Calculations What to Do If Nothing Works Summary 259 Questions and Problems 260 Chapter 8 Sampling Designs: Random, Adaptive, and Systematic Sampling Simple Random Sampling Estimates of Parameters Estimation of a Ratio Proportions and Percentages Statified Random Sampling Estimates of Parameters Allocation of Sample Size Construction of Strata Proportions and Percentages Adaptive Sampling Adaptive Cluster Sampling Statified Adaptive Cluster Sampling Systematic Sampling Multistage Sampling Sampling Units of Equal Size Sampling Units of Unequal Size 298

5 viii Contents 8.6 Summary 299 Questions and Problems,,301 Chapter 9 Sequential Sampling Two Alternative Hypotheses Means from a Normal Distribution Variances from a Normal Distribution Proportions from a Binomial Distribution ^4 Counts from a Negative Binomial Distribution Three Alternative Hypotheses Stopping Rules Kuno's Stopping Rule Green's Stopping Rule Ecological Measurements Sequential Schnabel Estimation of Population Size Sampling Plans for Count Data General Models for Two Alternative Hypotheses from Quadrat Counts Validating Sequential Sampling Plans Summary 337 Questions and Problems 338 Chapter 10 Experimental Designs General Principles of Experimental Design Randomization Replication and Pseudoreplication Balancing and Blocking Types of Experiemental Designs Linear Additive Models Factorial Designs Randomized Block Designs Nested Designs Latin Square Designs Repeated Measure Designs Environmental Impact Studies Types of Disturbances Transient Response Studies Variability of Measurements Where Should I Go Next? Summary 369 Questions and Problems 370 Part Four Estimating Community Parameters 373 Chapter 11 Similarity Coefficients and Cluster Analysis Measurement of Similarity Binary Coefficients Distance Coefficients Correlation Coefficients Other Similarity Measures 387

6 Contents ", ix 11.2 Data Standardization Cluster Analysis Single Linkage Clustering Complete Linkage Clustering Average Linkage Clustering Recommendations for Classifications Other Multivariate Techniques, Direct Gradient Analysis Ordination Classification Summary 405 Questions and Problems 406 Chapter 12 Species Diversity Measures Background Problems Concepts of Species Diversity Species Richness Heterogeneity Evenness Species Richness Measures Rarefaction Method Jackknife Estimate Bootstrap Procedure t Species-Area Curve Estimates Heterogenity Measures Logarithmic Series Lognormal Distribution ' Simpson's Index Shannon-Wiener Function Brillouin Index Evenness Measures Recommendations Summary '451 Questions and Problems 452 Chapter 13 Niche Measures and Resource Preferences What Is a Resource? Niche Breadth Levins's Measure Shannon-Wiener Measure Smith's Measure Number of Frequently Used Resources Niche Overlap MacArthur and Levins's Measure Percentage Overlap Morisita's Measure Simplified Morisita Index Horn's Index Hurlbert's Index Which Overlap Index Is Best? 472

7 x Contents 13.4 Measurement of Habitat and Dietary Preferences Forage Ratio Murdoch's Index Manly'sa ' Rank Preference Index Rodgers's Index for Cafeteria Experiments Which Preference Index? Summary J 492 Questions and Problems 493 Part Five Ecological Miscellanea 497 Chapter 14 Estimation of Survival Rates Finite and Instantaneous Rates Estimation from Life Tables Methods of Collecting Life Table Data Key Factor Analysis Expectation of Further Life Estimation of Survival from Age Composition Radiotelemetry Estimates of Survival Maximum Likelihood Method Kaplan-Meier Method Estimation of Bird Survival Rates Testing for Differences in Survival Rates Log-Rank Test Likelihood Ratio Test Temporal Differences in Mortality Rates Summary 538 Questions and Problems 539 Chapter 15 The Garbage Can Transformations Standard Transformations Box-Cox Transformation Repeatability Central Trend Lines in Regression Measuring Temporal Variability of Populations Jackknife and Bootstrap Techniques Summary 572 Questions and Problems 573 Appendices 577 References 581 Index 607

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