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A Primer of Ecological Statistics
SECOND EDITION
This page intentionally left blank
A Primer of
Ecological Statistics Second Edition

Nicholas J. Gotelli University of Vermont

Aaron M. Ellison Harvard Forest

Sinauer Associates, Inc. Publishers

Sunderland, Massachusetts U.S.A.
Cover art copyright © 2004 Elizabeth Farnsworth. See pages xx–xxi.

A PRIMER OF ECOLOGICAL STATISTICS, SECOND EDITION

Copyright © 2013 by Sinauer Associates, Inc. All rights reserved.
This book may not be reproduced without permission of the publisher.

For information address Sinauer Associates Inc., 23 Plumtree Road,

Sunderland, MA 01375 U.S.A.
FAX 413-549-1118
EMAIL [email protected], [email protected]
WEBSITE www.sinauer.com

Library of Congress Cataloging-in-Publication Data

Gotelli, Nicholas J., 1959-
A primer of ecological statistics / Nicholas J. Gotelli, University of Vermont, Aaron
M. Ellison, Harvard University. -- Second edition.
pages ; cm
Includes bibliographical references and index.
ISBN 978-1-60535-064-6
1. Ecology--Statistical methods. I. Ellison, Aaron M., 1960- II. Title.
QH541.15.S72G68 2013
577.072--dc23
2012037594

Printed in U.S.A.
5 4 3 2 1
 For
Maryanne
&
Elizabeth
Who measures heaven, earth, sea, and sky
Thus seeking lore or gaiety
Let him beware a fool to be.
SEBASTIAN BRANT, Ship of Fools, 1494. Basel, Switzerland.

[N]umbers are the words without which exact description of any natural
phenomenon is impossible…. Assuredly, every objective phenomenon, of
whatever kind, is quantitative as well as qualitative; and to ignore the for-
mer, or to brush it aside as inconsequential, virtually replaces objective
nature by abstract toys wholly devoid of dimensions — toys that neither
exist nor can be conceived to exist.
E. L. MICHAEL, “Marine ecology and the coefficient of association: A plea in
behalf of quantitative biology,” 1920. Journal of Ecology 8: 54–59.

[W]e now know that what we term natural laws are merely statistical
truths and thus must necessarily allow for exceptions. …[W]e need the
laboratory with its incisive restrictions in order to demonstrate the invari-
able validity of natural law. If we leave things to nature, we see a very dif-
ferent picture: every process is partially or totally interfered with by
chance, so much so that under natural circumstances a course of events
absolutely conforming to specific laws is almost an exception.
CARL JUNG, Foreword to The I Ching or Book of Changes.Third Edition,
1950, translated by R. Wilhelm and C. F. Baynes. Bollingen Series XIX,
Princeton University Press.
Brief Contents

PART I
FUNDAMENTALS OF PROBABILITY AND
STATISTICAL THINKING
1 An Introduction to Probability 3
2 Random Variables and Probability Distributions 25
3 Summary Statistics: Measures of Location and Spread 57
4 Framing and Testing Hypotheses 79
5 Three Frameworks for Statistical Analysis 107
PART II
DESIGNING EXPERIMENTS
6 Designing Successful Field Studies 137
7 A Bestiary of Experimental and Sampling Designs 163
8 Managing and Curating Data 207
PART III
DATA ANALYSIS
9 Regression 239
10 The Analysis of Variance 289
11 The Analysis of Categorical Data 349
12 The Analysis of Multivariate Data 383
PART IV
ESTIMATION
13 The Measurement of Biodiversity 449
14 Detecting Populations and Estimating their Size 483
Appendix Matrix Algebra for Ecologists 523
Contents

PART I
Fundamentals of
Probability and
Statistical Thinking
CHAPTER 1
An Introduction to
Probability 3 Bayes’ Theorem 22
What Is Probability? 4 Summary 24
Measuring Probability 4
The Probability of a Single Event: CHAPTER 2
Prey Capture by Carnivorous Plants 4
Estimating Probabilities by Sampling 7
Random Variables and
Problems in the Definition of Probability 9 Probability Distributions 25
The Mathematics of Probability 11 Discrete Random Variables 26
Defining the Sample Space 11 Bernoulli Random Variables 26
Complex and Shared Events: Combining An Example of a Bernoulli Trial 27
Simple Probabilities 13 Many Bernoulli Trials = A Binomial Random
Probability Calculations: Milkweeds and Variable 28
Caterpillars 15 The Binomial Distribution 31
Complex and Shared Events: Rules for Poisson Random Variables 34
Combining Sets 18
An Example of a Poisson Random Variable:
Conditional Probabilities 21 Distribution of a Rare Plant 36
 Contents ix

The Expected Value of a Discrete Random CHAPTER 4

Variable 39
The Variance of a Discrete Random Variable 39 Framing and Testing
Continuous Random Variables 41 Hypotheses 79
Uniform Random Variables 42 Scientific Methods 80
The Expected Value of a Continuous Random Deduction and Induction 81
Variable 45
Modern-Day Induction: Bayesian Inference 84
Normal Random Variables 46
The Hypothetico-Deductive Method 87
Useful Properties of the Normal
Distribution 48 Testing Statistical Hypotheses 90
Other Continuous Random Variables 50 Statistical Hypotheses versus Scientific
Hypotheses 90
The Central Limit Theorem 53
Statistical Significance and P-Values 91
Summary 54 Errors in Hypothesis Testing 100
Parameter Estimation and Prediction 104
CHAPTER 3 Summary 105
Summary Statistics: Measures
of Location and Spread 57 CHAPTER 5
Measures of Location 58 Three Frameworks for
The Arithmetic Mean 58 Statistical Analysis 107
Other Means 60 Sample Problem 107
Other Measures of Location: The Median and
Monte Carlo Analysis 109
the Mode 64
Step 1: Specifying the Test Statistic 111
When to Use Each Measure of Location 65
Step 2: Creating the Null Distribution 111
Measures of Spread 66
Step 3: Deciding on a One- or Two-Tailed
The Variance and the Standard Deviation 66 Test 112
The Standard Error of the Mean 67 Step 4: Calculating the Tail Probability 114
Skewness, Kurtosis, and Central Moments 69 Assumptions of the Monte Carlo Method 115
Quantiles 71 Advantages and Disadvantages of the Monte
Using Measures of Spread 72 Carlo Method 115
Some Philosophical Issues Surrounding Parametric Analysis 117
Summary Statistics 73 Step 1: Specifying the Test Statistic 117
Confidence Intervals 74 Step 2: Specifying the Null Distribution 119
Generalized Confidence Intervals 76 Step 3: Calculating the Tail Probability 119
Summary 78 Assumptions of the Parametric Method 120
Advantages and Disadvantages of the
Parametric Method 121
x Contents

Non-Parametric Analysis: A Special Case of Step 4: Calculating the Likelihood 129

Monte Carlo Analysis 121 Step 5: Calculating the Posterior Probability
Bayesian Analysis 122 Distribution 129
Step 1: Specifying the Hypothesis 122 Step 6: Interpreting the Results 130
Step 2: Specifying Parameters as Random Assumptions of Bayesian Analysis 132
Variables 125 Advantages and Disadvantages of Bayesian
Step 3: Specifying the Prior Probability Analysis 133
Distribution 125 Summary 133

PART II
Designing Experiments
CHAPTER 6
Designing Successful
Field Studies 137
What Is the Point of the Study? 137
Are There Spatial or Temporal Differences in
Variable Y? 137
What Is the Effect of Factor X on Large-Scale Studies and Environmental
Variable Y? 138 Impacts 150
Are the Measurements of Variable Y Consistent Ensuring Independence 151
with the Predictions of Hypothesis H? 138
Avoiding Confounding Factors 153
Using the Measurements of Variable Y,
What Is the Best Estimate of Parameter θ Replication and Randomization 154
in Model Z? 139 Designing Effective Field Experiments and
Manipulative Experiments 139 Sampling Studies 158
Natural Experiments 141 Are the Plots or Enclosures Large Enough to
Ensure Realistic Results? 158
Snapshot versus Trajectory Experiments 143
What Is the Grain and Extent of the Study? 158
The Problem of Temporal Dependence 144
Does the Range of Treatments or Census
Press versus Pulse Experiments 146 Categories Bracket or Span the Range of
Replication 148 Possible Environmental Conditions? 159
How Much Replication? 148 Have Appropriate Controls Been Established
to Ensure that Results Reflect Variation Only
How Many Total Replicates Are Affordable? 149 in the Factor of Interest? 160
The Rule of 10 150
 Contents xi

Have All Replicates Been Manipulated in the CHAPTER 8

Same Way Except for the Intended
Treatment Application? 160 Managing and Curating
Have Appropriate Covariates Been Measured
in Each Replicate? 161 Data 207
Summary 161 The First Step: Managing Raw Data 208
Spreadsheets 208
CHAPTER 7 Metadata 209
The Second Step: Storing and Curating the
A Bestiary of Experimental Data 210
and Sampling Designs 163 Storage: Temporary and Archival 210
Categorical versus Continuous Variables 164 Curating the Data 211
Dependent and Independent Variables 165 The Third Step: Checking the Data 212
Four Classes of Experimental Design 165 The Importance of Outliers 212
Regression Designs 166 Errors 214
ANOVA Designs 171 Missing Data 215
Alternatives to ANOVA: Experimental Detecting Outliers and Errors 215
Regression 197 Creating an Audit Trail 223
Tabular Designs 200 The Final Step: Transforming the Data 223
Alternatives to Tabular Designs: Proportional Data Transformations as a Cognitive Tool 224
Designs 203 Data Transformations because the Statistics
Summary 204 Demand It 229
Reporting Results: Transformed or Not? 233
The Audit Trail Redux 233
Summary: The Data Management Flow
Chart 235

PART III
Data Analysis
CHAPTER 9
Regression 239
Defining the Straight Line and Its Two
Parameters 239
Fitting Data to a Linear Model 241
Variances and Covariances 244
xii Contents

Least-Squares Parameter Estimates 246 Hypothesis Tests with ANOVA 296

Variance Components and the Coefficient of Constructing F-Ratios 298
Determination 248 A Bestiary of ANOVA Tables 300
Hypothesis Tests with Regression 250 Randomized Block 300
The Anatomy of an ANOVA Table 251 Nested ANOVA 302
Other Tests and Confidence Intervals 253 Two-Way ANOVA 304
Assumptions of Regression 257 ANOVA for Three-Way and n-Way Designs 308
Diagnostic Tests For Regression 259 Split-Plot ANOVA 308
Plotting Residuals 259 Repeated Measures ANOVA 309
Other Diagnostic Plots 262 ANCOVA 314
The Influence Function 262 Random versus Fixed Factors in ANOVA 317
Monte Carlo and Bayesian Analyses 264 Partitioning the Variance in ANOVA 322
Linear Regression Using Monte Carlo After ANOVA: Plotting and Understanding
Methods 264 Interaction Terms 325
Linear Regression Using Bayesian Methods 266 Plotting Results from One-Way ANOVAs 325
Other Kinds of Regression Analyses 268 Plotting Results from Two-Way ANOVAs 327
Robust Regression 268 Understanding the Interaction Term 331
Quantile Regression 271 Plotting Results from ANCOVAs 333
Logistic Regression 273 Comparing Means 335
Non-Linear Regression 275 A Posteriori Comparisons 337
Multiple Regression 275 A Priori Contrasts 339
Path Analysis 279 Bonferroni Corrections and the Problem of
Model Selection Criteria 282 Multiple Tests 345
Model Selection Methods for Multiple Summary 348
Regression 283
Model Selection Methods in Path Analysis 284
CHAPTER 11
Bayesian Model Selection 285
Summary 287 The Analysis of Categorical
Data 349
CHAPTER 10 Two-Way Contingency Tables 350
The Analysis of Variance 289 Organizing the Data 350
Are the Variables Independent? 352
Symbols and Labels in ANOVA 290
Testing the Hypothesis: Pearson’s Chi-square
ANOVA and Partitioning of the Sum of Test 354
Squares 290 An Alternative to Pearson’s Chi-Square:
The Assumptions of ANOVA 295 The G-Test 358
 Contents xiii

The Chi-square Test and the G-Test for R × C The Multivariate Normal Distribution 394
Tables 359 Testing for Multivariate Normality 396
Which Test To Choose? 363
Measurements of Multivariate Distance 398
Multi-Way Contingency Tables 364 Measuring Distances between Two
Organizing the Data 364 Individuals 398
On to Multi-Way Tables! 368 Measuring Distances between Two Groups 402
Bayesian Approaches to Contingency Tables 375 Other Measurements of Distance 402
Tests for Goodness-of-Fit 376 Ordination 406
Goodness-of-Fit Tests for Discrete Principal Component Analysis 406
Distributions 376 Factor Analysis 415
Testing Goodness-of-Fit for Continuous Principal Coordinates Analysis 418
Distributions: The Kolmogorov-Smirnov
Test 380 Correspondence Analysis 421
Summary 382 Non-Metric Multidimensional Scaling 425
Advantages and Disadvantages of Ordination
427
CHAPTER 12
Classification 429
The Analysis of Multivariate Cluster Analysis 429
Data 383 Choosing a Clustering Method 430
Discriminant Analysis 433
Approaching Multivariate Data 383
Advantages and Disadvantages of
The Need for Matrix Algebra 384 Classification 437
Comparing Multivariate Means 387 Multivariate Multiple Regression 438
Comparing Multivariate Means of Two Redundancy Analysis 438
Samples: Hotelling’s T 2 Test 387
Summary 444
Comparing Multivariate Means of More Than
Two Samples: A Simple MANOVA 390

PART IV
Estimation
CHAPTER 13
The Measurement of Biodiversity
449
Estimating Species Richness 450
Standardizing Diversity Comparisons through
Random Subsampling 453
xiv Contents

Rarefaction Curves: Interpolating Species Occupancy of More than One Species 493
Richness 455 A Hierarchical Model for Parameter Estima-
The Expectation of the Individual-Based Rar- tion and Modeling 495
efaction Curve 459 Occupancy Models for Open Populations 501
Sample-Based Rarefaction Curves: Massachu- Dynamic Occupancy of the Adelgid in Massa-
setts Ants 461 chusetts 505
Species Richness versus Species Density 465 Estimating Population Size 506
The Statistical Comparison of Rarefaction Mark-Recapture: The Basic Model 507
Curves 466
Mark-Recapture Models for Open Populations
Assumptions of Rarefaction 467 516
Asymptotic Estimators: Extrapolating Occupancy Modeling and Mark-Recapture:
Species Richness 470 Yet More Models 518
Rarefaction Curves Redux: Extrapolation and Sampling for Occupancy and Abundance
Interpolation 476 519
Estimating Species Diversity and Evenness Software for Estimating Occupancy and
476 Abundance 521
Hill Numbers 479
Summary 522
Software for Estimation of Species Diversity
481
APPENDIX
Summary 482
Matrix Algebra for Ecologists 523
CHAPTER 14
Glossary 535
Detecting Populations
and Estimating their Size 483 Literature Cited 565
Occupancy 485 Index 583
The Basic Model: One Species, One Season,
Two Samples at a Range of Sites 487
Preface

What is the “added value” for readers in purchasing this Second Edition? For this
Second Edition, we have added a new Part 4, on “Estimation,” which includes
two chapters: “The Measurement of Biodiversity” (Chapter 13), and “Detect-
ing Populations and Estimating their Size” (Chapter 14). These two new chap-
ters describe measurements and methods that are beginning to be widely used
and that address questions that are central to the disciplines of community ecol-
ogy and demography. Some of the methods we describe have been developed
only in the last 10 years, and many of them continue to evolve rapidly.
The title of this new section, “Estimation,” reflects an ongoing shift in ecolo-
gy from testing hypotheses to estimating parameters. Although Chapter 3
describes summary measures of location and spread, most of the focus of the
First Edition—and of many ecological publications—is on hypothesis testing
(Chapter 4). In the two new chapters, we are more concerned with the process
of estimation itself, although the resulting estimators of biodiversity and pop-
ulation size certainly can be used in conventional statistical tests once they are
calculated. It is useful to think of biodiversity and population sizes as latent or
unknown “state variables.” We want to measure these things, but it is impossi-
ble to find and tag all of the trout in a stream or discover all of the species of ants
that lurk in a bog. Instead, we work only with small samples of diversity, or lim-
ited collections of marked animals, and we use these samples to estimate the
“true” value of the underlying state variables. The two new chapters are united
not only by their focus on estimation, but also by their common underlying sta-
tistical framework: methods for asymptotic estimators of species richness dis-
cussed in Chapter 13 were derived from methods developed for mark-recapture
studies discussed in Chapter 14. We hope the methods described in these two
chapters will provide some fresh statistical tools for ecologists to complement
the more classical topics we cover in Chapters 1–12.
Chapter 13 begins with what seems like a relatively simple question: how
many species are there in an assemblage? The central problem is that nature is
xvi Preface

(still) vast, our sample of its diversity is tiny, and there are always many rare and
undetected species. Moreover, because the number of species we count is heav-
ily influenced by the number of individuals and samples we have collected, the
data must be standardized for meaningful comparisons. We describe methods
for both interpolation (rarefaction) and extrapolation (asymptotic species rich-
ness estimators) to effectively standardize biodiversity comparisons. We also
advocate the use of Hill numbers as a class of diversity indices that have useful
statistical properties, although they too are subject to sampling effects. We thank
Anne Chao, Rob Colwell, and Lou Jost for their ongoing collaboration, as well
as their insights, extended correspondence, and important contributions to the
literature on biodiversity estimation.
In Chapter 14, we delve deeper into the problem of incomplete detection,
addressing two main questions. First, how do we estimate the probability that
we can detect a species, given that it is actually present at a site? Second, for a
species that is present at a site, how large is its population? Because we have a
limited sample of individuals that we can see and count, we must first estimate
the detection probability of an individual species before we can estimate the size
of its population. Estimates of occupancy probability and population size are
used in quantitative models for the management of populations of both rare
and exploited species. And, just as we use species richness estimators to extrap-
olate beyond our sample, we use estimates from occupancy models and mark-
recapture studies to extrapolate and forecast what will happen to these popula-
tions as, for example, habitats are fragmented, fishing pressure increases, or the
climate changes. We thank Elizabeth Crone, Dave Orwig, Evan Preisser, and Rui
Zhang for letting us use some of their unpublished data in this chapter and for
discussing their analyses with us. Bob Dorazio and Andy Royle discussed these
models with us; they, along with Evan Cooch, have developed state-of-the-art
open-source software for occupancy modeling and mark-recapture analysis.
In this edition, we have also corrected numerous minor errors that astute read-
ers have pointed out to us over the past 8 years. We are especially grateful for the
careful reading of Victor Lemes Landeiro, who, together with his colleagues Fabri-
cio Beggiato Baccaro, Helder Mateus Viana Espirito Santo, Miriam Plaza Pinto,
and Murilo Sversut Dias, translated the entire book into Portuguese. Specific con-
tributions of these individuals, and all the others who sent us comments and
pointed out errors, are noted in the Errata section of the book’s website (har-
vardforest.fas.harvard.edu/ellison/publications/primer/errata). Although we have
corrected all errors identified in the first edition, others undoubtedly remain
(another problem in detection probability); please contact us if you find any.
All code posted on the Data and Code section of the book’s website (har-
vardforest.fas.harvard.edu/ellison/publications/primer/datafiles) that we used
 Preface xvii

for the analyses in the book has been updated and reworked in the R program-
ming language (r-project.org). In the time since we wrote the first edition, R has
become the de facto standard software for statistical analysis and is used by most
ecologists and statisticians for their day-to-day analytical work. We have pro-
vided some R scripts for our figures and analyses, but they have minimal anno-
tation, and there are already plenty of excellent resources available for ecologi-
cal modeling and graphics with R.
Last but not least, we thank Andy Sinauer, Azelie Aquadro, Joan Gemme, Chris
Small, Randy Burgess, and the entire staff at Sinauer Associates for carefully edit-
ing our manuscript pages and turning them into an attractive book.1

Preface to the First Edition

Why another book on statistics? The field is already crowded with texts on bio-
metry, probability theory, experimental design, and analysis of variance. The
short answer is that we have yet to find a single text that meets the two specific
needs of ecologists and environmental scientists. The first need is a general intro-
duction to probability theory including the assumptions of statistical inference
and hypothesis testing. The second need is a detailed discussion of specific
designs and analyses that are typically encountered in ecology.
This book reflects several collective decades of teaching statistics to under-
graduate and graduate students at the University of Oklahoma, the University
of Vermont, Mount Holyoke College, and the Harvard Forest. The book repre-
sents our personal perspective on what is important at the interface of ecology
and statistics and the most effective way to teach it. It is the book we both wish
we had in our hands when we started studying ecology in college.
A “primer” suggests a short text, with simple messages and safe recommen-
dations for users. If only statistics were so simple! The subject is vast, and new
methods and software tools for ecologists appear almost monthly. In spite of the
book’s length (which stunned even us), we still view it as a distillation of statis-
tical knowledge that is important for ecologists. We have included both classic
expositions as well as discussions of more recent methods and techniques. We
hope there will be material that will appeal to both the neophyte and the expe-
rienced researcher. As in art and music, there is a strong element of style and
personal preference in the choice and application of statistics. We have explained
our own preferences for certain kinds of analyses and designs, but readers and
users of statistics will ultimately have to choose their own set of statistical tools.

1
And for dealing with all of the footnotes!
xviii Preface

How To Use This Book

In contrast to many statistics books aimed at biologists or ecologists—“biome-
try” texts—this is not a book of “recipes,” nor is it a set of exercises or problems
tied to a particular software package that is probably already out of date.
Although the book contains equations and derivations, it is not a formal statis-
tical text with detailed mathematical proofs. Instead, it is an exposition of basic
and advanced topics in statistics that are important specifically for ecologists
and environmental scientists. It is a book that is meant to be read and used, per-
haps as a supplement to a more traditional text, or as a stand-alone text for
students who have had at least a minimal introduction to statistics. We hope this
book will also find a place on the shelf (or floor) of environmental profession-
als who need to use and interpret statistics daily but who may not have had
formal training in statistics or have ready access to helpful statisticians.
Throughout the book we make extensive use of footnotes. Footnotes give us
a chance to greatly expand the material and to talk about more complex issues
that would bog down the flow of the main text. Some footnotes are purely his-
torical, others cover mathematical and statistical proofs or details, and others
are brief essays on topics in the ecological literature. As undergraduates, we both
were independently enamored of Hutchinson’s classic An Introduction to Popu-
lation Biology (Yale University Press 1977). Hutchinson’s liberal use of footnotes
and his frequent forays into history and philosophy served as our stylistic model.
We have tried to strike a balance between being concise and being thor-
ough. Many topics recur in different chapters, although sometimes in slightly
different contexts. Figure and table legends are somewhat expansive because they
are meant to be readable without reference to the text. Because Chapter 12
requires matrix algebra, we provide a brief Appendix that covers the essentials
of matrix notation and manipulation. Finally, we have included a comprehen-
sive glossary of terms used in statistics and probability theory. This glossary may
be useful not only for reading this book, but also for deciphering statistical meth-
ods presented in journal articles.

Mathematical Content
Although we have not shied away from equations when they are needed, there
is considerably less math here than in many intermediate level texts. We also try
to illustrate all methods with empirical analyses. In almost all cases, we have used
data from our own studies so that we can illustrate the progression from raw
data to statistical output.
Although the chapters are meant to stand alone, there is considerable cross-
referencing. Chapters 9–12 cover the traditional topics of many biometry texts
and contain the heaviest concentration of formulas and equations. Chapter 12,
 Preface xix

on multivariate methods, is the most advanced in the book. Although we have

tried to explain multivariate methods in plain English (no easy task!), there is
no way to sugar-coat the heavy dose of matrix algebra and new vocabulary nec-
essary for multivariate analysis.

Coverage and Topics

Statistics is an evolving field, and new methods are always being developed to
answer ecological questions. This text, however, covers core material that we
believe is foundational for any statistical analysis. The book is organized around
design-based statistics—statistics that are to be used with designed observational
and experimental studies. An alternative framework is model-based statistics,
including model selection criteria, likelihood analysis, inverse modeling, and
other such methods.
What characterizes the difference between these two approaches? In brief,
design-based analyses primarily address the problem of P(data | model): assess-
ing the probability of the data given a model that we have specified. An impor-
tant element of design-based analysis is being explicit about the underlying
model. In contrast, model-based statistics address the problem of P (model |
data)—that is, assessing the probability of a model given the data that are pres-
ent. Model-based methods are often most appropriate for large datasets that may
not have been collected according to an explicit sampling strategy.
We also discuss Bayesian methods, which somewhat straddle the line between
these two approaches. Bayesian methods address P(model | data), but can be
used with data from designed experiments and observational studies.
The book is divided into three parts. Part I discusses the fundamentals of
probability and statistical thinking. It introduces the logic and language of prob-
ability (Chapter 1), explains common statistical distributions used in ecology
(Chapter 2), and introduces important measures of location and spread (Chap-
ter 3). Chapter 4 is more of a philosophical discussion about hypothesis testing,
with careful explanations of Type I and Type II errors and of the ubiquitous
statement that “P < 0.05.” Chapter 5 closes this section by introducing the three
major paradigms of statistical analysis (frequentist, Monte Carlo, and Bayesian),
illustrating their use through the analysis of a single dataset.
Part II discusses how to successfully design and execute observational studies
and field experiments. These chapters contain advice and information that is to
be used before any data are collected in the field. Chapter 6 addresses the practi-
cal problems of articulating the reason for the study, setting the sample size, and
dealing with independence, replication, randomization, impact studies, and envi-
ronmental heterogeneity. Chapter 7 is a bestiary of experimental designs. How-
ever, in contrast to almost all other texts, there are almost no equations in this
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