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Samuel M. Scheiner

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This page intentionally left blank
Design and Analysis of
Ecological Experiments

SECOND EDITION

Edited by
Samuel M. Scheiner and
Jessica Gurevitch

OXFORD
UNIVERSITY PRESS
2001
OXFORD
UNIVERSITY PRESS
Oxford New York
Athens Auckland Bangkok Bogota Buenos Aires Calcutta
Cape Town Chennai Dar es Salaam Delhi Florence Hong Kong Istanbul
Karachi Kuala Lumpur Madrid Melbourne Mexico City Mumbai
Nairobi Paris Sao Paulo Shanghai Singapore Taipei Tokyo Toronto Warsaw
and associated companies in
Berlin Ibadan

Copyright © 2001 by Oxford University Press, Inc.

Published by Oxford University Press, Inc.
198 Madison Avenue, New York, New York 10016
Oxford is a registered trademark of Oxford University Press
All rights reserved. No part of this publication may be reproduced,
stored in a retrieval system, or transmitted, in any form or by any means,
electronic, mechanical, photocopying, recording, or otherwise,
without the prior permission of the Oxford University Press.

Library of Congress Cataloging-in-Publication Data

Design and analysis of ecological experiments / edited by Samuel M. Scheiner and
Jessica Gurevitch.—2nd ed.
p. cm.
Includes bibliographical references.
ISBN 0-19-513187-8; 0-19-513188-6 (pbk.)
1. Ecology—Statistical methods. 2. Experimental design. I. Scheiner, Samuel M..
1956- II. Gurevitch, Jessica.
QH541.15.S72 D47 2000
577'.07'27-dc21 00-035647

9 8 7 6 5 4 3 2 1
Printed in the United States of America
on recycled acid-free paper
For Edith, Judy, and Kayla
and the memory of
my father, Sayre
S.M.S.

Dedicated with much appreciation to

my father, Louis Gurevitch,
and in loving memory of
my mother, Esther Gurevitch
J.G.

We note with deep sadness

the passing of one
of our contributors,
Tom Frost.
This page intentionally left blank
 Preface to the Second Edition

This second edition of our book on advanced statistical techniques for

ecologists comes out seven years after the first. We are very pleased with the
reception of the first edition, which has been widely read and used in many
graduate seminars on statistics in ecological research. The enthusiasm expressed
for the book was part of the motivation for this new edition, which updates and
expands on the original. We like to think that this book has played a role in
introducing many previously unfamiliar approaches to ecologists and in setting
higher standards for the use and reporting of more conventional analyses.
This new edition is designed to help continue and propel these trends. Nearly
all of the chapters included from the first edition are revised, several substantially,
to bring them up to date. We added four new chapters that reflect recent advances
in statistical practice, particularly within ecology. The changes result from a com-
bination of new statistical theory, new computer software and hardware capabili-
ties, and evolving practices and improved statistical standards among ecologists.
We are also taking advantage of another important advance in information
technology, the World Wide Web. Oxford University Press has established a
companion Website [http://www.oup-usa.org/sc/0195131878/] that contains the
source code and data sets for many of the chapters. By moving the source code
to a Website, we were able to add a chapter to the book. Readers can now check
their results against the sample data sets on-line.
It is exciting to be publishing a book in 2001. Ecological research and the use
of statistics in ecology continue to evolve rapidly. We hope that this new edition
will prove helpful to established researchers and to those in training, who will
viii Design and Analysis of Ecological Experiments

bring new standards and new insights into the service of the science in the twenty-
first century.
Samuel M. Scheiner
Jessica Gurevitch
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 Preface to the First Edition

The genesis of this book was the result of a conversation while eating a
bento (a Japanese box lunch of rice, sushi, pickles, and other delicacies) on a
sidewalk in Yokohama in 1990. The conversation had turned to statistics and
both of us were commenting on statistical issues and techniques that were either
underused or misused by ecologists. For some time, Jessica had been contemplat-
ing a book on statistical techniques in experimental design and analysis, written
for ecologists. The goal of such a book was both to encourage the correct use of
some of the more well-known approaches, and to make some potentially very
useful but less well known techniques available to ecologists. We both felt
strongly that such a book was timely, and would be useful to ecologists working
on applied as well as basic problems. Had Sam not intervened, this idea would
undoubtedly have met the fate of many other fine ideas that have never made it
into the light of day.
It was apparent to both of us that we did not have the skills to write such a
book ourselves, nor the time. However, we were able to compile a list of topics
and potential contributors whom we knew were knowledgeable about those top-
ics. (An initial outline for the book was composed while sitting for hours, stalled
in traffic going to and from Mount Fuji. At the next INTECOL meeting keep
your eyes out for "I survived the trip to Mt. Fuji" t-shirts.) We batted (actually
e-mailed) these ideas back and forth for nearly a year. The success of a sympo-
sium organized by Phil Dixon at the 1991 annual meeting of the Ecological Soci-
ety of America on the design of ecological experiments encouraged us to continue
our endeavors, and we managed to secure commitments from many of our con-
tributors during that week. The enthusiasm for this undertaking expressed by
x Design and Analysis of Ecological Experiments

colleagues we spoke with buoyed us, as did the interest and encouragement of
Greg Payne from Chapman and Hall. Therefore, despite warnings about the tra-
vails of editing a book, we forged ahead. So—beware of the dangers of conversa-
tion over raw fish.
Samuel M. Schemer
Jessica Gurevitch
 Acknowledgments

We wish to thank a number of people who contributed to the produc-

tion of this volume. We are grateful to those colleagues who painstakingly re-
viewed chapters: Marti Anderson, Norma Fowler, Charles Goodnight, Ed Green,
Jeff Hatfield, Ray Hilborn, Ed Heske, Charles Janson, Martin Lechowicz, Bryan
Manly, Jim McGraw, Tom Meagher, Scott Menard, Tom Mitchell-Olds, Patrick
Phillips, Alan Tessier, Joel Trexler, Art Weis, and Neil Willits. We offer special
thanks to Greg Payne, book editor of the first edition, and Kirk Jensen, book
editor of the second edition, for their efforts. A fellowship from the Arnold Arbo-
retum of Harvard University, responsible for sending J.G. to Japan and thus for
the inception of this book, is gratefully acknowledged. S.M.S. was supported for
his trip to Japan by funds from the Department of Biological Sciences, the Col-
lege of Liberal Arts & Sciences and the Graduate School of Northern Illinois
University, and the City of Yokohama; the Department of Biological Sciences
provided support during the editorial process. S.M.S. thanks Judy Scheiner for all
of her patience and forbearance during the production of this book. J.G. extends
many thanks to Todd Postol for being in her corner all of these years.
This page intentionally left blank
Contents

Contributors xv

1 Theories, Hypotheses, and Statistics 3

Samuel M. Scheiner

2 Power Analysis and Experimental Design 14

Robert J. Steidl and Len Thomas

3 Exploratory Data Analysis and Graphic Display 37

Aaron M. Ellison

4 ANOVA: Experimental Layout and Analysis 63

Catherine Potvin

5 ANOVA and ANCOVA: Field Competition Experiments 77

Deborah E. Goldberg and Samuel M. Scheiner

6 MANOVA: Multiple Response Variables and

Multispecies Interactions 99
Samuel M. Scheiner

7 ANCOVA: Nonparametric and Randomization

Approaches I 16
Peter S. Petraitis, Steven J. Beaupre, and Arthur E. Dunham
xiv Design and Analysis of Ecological Experiments

8 Repeated-measures Analysis: Growth and Other

Time-dependent Measures 134
Carl N. von Ende

9 Time Series Intervention Analysis: Unreplicated Large-scale

Experiments 158
Paul W. Rasmussen, Dennis M. Heisey, Erik V. Nordheim, and
Thomas M. Frost

10 Nonlinear Curve Fitting: Predation and Functional Response

Curves 178
Steven A. Juliano

II Logit Modeling and Logistic Regression: Aphids, Ants, and

Plants 197
Ted Floyd

12 Path Analysis: Pollination 217

Randall J. Mitchell

13 Failure-time Analysis: Studying Times to Events and Rates

at Which Events Occur 235
Gordon A. Fox

14 The Bootstrap and the Jackknife: Describing the Precision

of Ecological Indices 267
Philip M. Dixon

15 Spatial Statistics: Analysis of Field Experiments 289

Jay M. Ver Hoef and Noel Cressie

16 Mantel Tests: Spatial Structure in Field Experiments 308

Marie-Josee Fortin and Jessica Gurevitch

17 Bayesian Statistics: Estimating Plant Demographic

Parameters 327
James S. Clark and Michael Lavine

18 Meta-analysis: Combining the Results of Independent

Experiments 347
Jessica Gurevitch and Larry V. Hedges

References 371

Index 403
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