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On the Cover
Sandhill cranes (Crus canadensis) take off from their night refuge from
predators en route to nearby fields to forage for the day. Bosque del Apache
National Wildlife Refuge, New Mexico. Photograph© William D. Bowman.

Ecology, Fourth Edition

Copyright© 2017 by Sinauer Associates, Inc. All rights reserved.
This book may not be reproduced in whole or in part without permission.

Address inquiries and orders to:

Sinauer Associates, Inc., P.O. Box 407, Sunderland, MA 01375-0407 U.S.A.

www.sinauer.com

FAX: 413-549-1118
E-mail: [email protected]

Library of Congress Cataloging-in-Publication Data

Names: Bowman, William D. I Hacker, Sally D. I Cain, Michael L. (Michael

Lee), 1956-
Title: Ecology I William D. Bowman, University of Colorado, Sally D. Hacker,
Oregon State University, Michael L. Cain, New Mexico State University.
Description: Fourth edition. I Sunderland, Massachusetts : Sinauer
Associates, Inc., c2017. I Includes bibliographical references and index.
Identifiers: LCCN 2017002873 I ISBN 9781605356181
Subjects: LCSH: Ecology--Textbooks.
Classification: LCC QH541 .E31933 2017 I DOC 577--dc23
LC record available at https:/ /lccn.loc.gov /2017002873

Printed in U.S.A.
54321
 For fen, Gordon, and Miles and their unwavering support,
and to my students for pushing me as much as I pushed them.
WDB

For my family and my students, whose gift of time

has made all the difference.
SDH

For Debra and Hannah, with thanks and love.

MLC
About the Authors

William D. Bowman is a Profes-

sor at the University of Colorado at
Boulder, affiliated with the Depart-
ment of Ecology and Evolutionary
Biology, Mountain Research Station,
and the Institute of Arctic and Alpine
Research. He earned his Ph.D. from
Duke University. Dr. Bowman has
taught courses in introductory ecol-
ogy, plant ecology, plant-£oil interac-
tions, and ecosystems ecology, and
for over two decades he has directed
undergraduate summer field courses
and research programs. His research
focuses on the intersections of physi-
ological ecology, community dynam-
ics, and ecosystem function, particu-
larly in the context of environmental
change.

Sally D. Hacker is a Professor at

Oregon State University, Corvallis,
where she has been a faculty member
since 2004. She has taught courses
in introductory ecology, community William D. Bowman Sally D. Hacker Michael L. Cain
ecology, and marine biology. She is
particularly interested in promot-
ing active and experiential learning textbooks, Ecology and Life: The Sci- and the Rose-Hulman Institute of
for students interested in ecology ence of Biology (Sinauer), she is author Technology. In addition to his work
and field experiences. Dr. Hacker's or coauthor on numerous articles and on this book, Dr. Cain is a coauthor
research explores the structure, func- book chapters exploring themes in of Campbell's Biology (Eleventh Edi-
tion, and services of natural and community ecology and ecosystem tion) and Biology in Focus (Second
managed ecosystems under varying functions and services. Edition). He has instructed students
contexts of species interactions and across a wide range of subjects, in-
global change. She has conducted Michael L. Cain, having opted to cluding introductory biology, ecol-
research with plants and animals focus full-time on writing, is affiliat- ogy, field ecology, evolution, botany,
in rocky intertidal, estuarine, and ed with the Department of Biology at mathematical biology, and biostatis-
coastal dune ecosystems. Her work New Mexico State University. After tics. His research interests include:
has most recently focused on the receiving his Ph.D. in Ecology and plant ecology; long-distance disper-
protective role of coastal ecosystems Evolutionary Biology from Cornell sal; ecological and evolutionary dy-
in mitigating the vulnerability from University, he was a faculty mem- namics in hybrid zones; and search
climate change. In addition to the ber at New Mexico State University behavior in plants and animals.
Brief Contents

1 The Web of Life 2 UnitS Communities 355

Unit1 Organisms and Their 16 The Nature of Communities 356
Environment 21 17 Change in Communities 377
z The Physical Environment 22 18 Biogeography 399
3 The Biosphere 50 19 Species Diversity
in Communities 424
4 Coping with Environmental Variation:
Temperature and Water 84 Unit6 Ecosystems 447
5 Coping with Environmental Variation:
20 Production 448
Energy 109
21 Energy Flow and Food Webs 470
Unit2 Evolutionary Ecology 135
22 Nutrient Supply and Cycling 493
6 Evolution and Ecology 136
Unit7 Applied and Large-Scale
7 Life History 160
Ecology 517
8 Behavioral Ecology 182
23 Conservation Biology 518
Unit3 Populations 203 24 Landscape Ecology and Ecosystem
9 Population Distribution and Management 544
Abundance 204 25 Global Ecology 568
10 Population Growth
and Regulation 226
11 Population Dynamics 249

Unit4 Species Interactions 271

12 Predation 272
13 Parasitism 294
14 Competition 317
15 Mutualism and Commensalism 336
Preface

Ecology is at the heart of understanding our world-it more fully below) about an applied problem or interest-
serves as the glue that brings together information from ing bit of natural history. Once students are drawn in by
a multitude of very different scientific disciplines, and the Case Study, the "storyline" that it initiates is main-
it integrates this information in a way that informs us tained throughout the rest of the chapter. We use a nar-
about how nature works. As our environment continues rative writing style to link the sections of the chapter to
to change at an alarming rate it becomes increasingly one another, thus helping students keep the big picture
critical that we improve our understanding of the reper- in mind. In addition, the sections of the chapter are or-
cussions of climate change, non-sustainable extraction of ganized around a small number of Key Concepts (also
resources, the spread of invasive species, and pollution. described more thoroughly below) that were carefully
Stewardship of resources that humanity requires-food, selected to summarize current knowledge and provide
clean water, clean air, and many others-is best under- students with a clear overview of the subject at hand .
stood through the lens of ecological understanding. Similarly, when designing the art, pedagogy came first.
Advances in ecology occur regularly, facilitated by Many students are visual learners, so we worked very
technological and computational breakthroughs, as well hard to ensure that each figure "tells a story" that can be
as creative experimental research. This continued ad- understood on its own.
vancement, along with the diversity of subjects that form As another way to help us achieve our primary goal
its basis, makes ecology a potentially daunting and com- of teaching students, we followed a "less is more" philos-
plicated subject to learn and teach. Covering the breadth ophy. We were guided by the principle that if we covered
of ecology sufficiently in a textbook requires care so that less material, but presented it clearly and well, students
students are not overwhelmed with the amount of mate- would learn more. Hence, our chapters are relatively
rial and instructors have adequate material to effective- short and they are built around a small number of Key
ly engage students. With these challenges in mind, the Concepts (typically, three to five). We made these choices
overarching goal for the Fourth Edition of Ecology was to to prevent students from being overwhelmed by long, dif-
enhance the book as a learning tool for students and as fuse chapters, and to allow them to master the big ideas
a teaching tool for professors. To achieve these goals, the first. In addition, we put our "less is more" philosophy
book's two core principles guided our every step. into action by asking each other whether the text served
one of the following purposes:
Core Principles Guiding Ecology, • Does it help to explain an essential concept?
Fourth Edition • Does it show how the process of ecological inquiry
works?
This book is written for undergraduate students taking
their first course in ecology. We set out to introduce our • Does it motivate readers by focusing on a key eco-
readers to the beauty and importance of ecology, and to logical application or a fascinating piece of natural
do so without presenting them with too much material or history?
boring them with unnecessary detail. While writing the This approach made for some tough choices, but it en-
Fourth Edition of Ecology, we kept our focus on two core abled us to focus on teaching students what is currently
principles: "Teaching Comes First" and "Less Is More." known about ecology without burdening them with ex-
Enabling effective teaching is our primary goal and cessive detail.
motivation in Ecology. The structure and content of our We also recognize that many instructors are choosing
chapters are designed primarily to make them good tools to "flip" their instructional style, with an emphasis on
for teaching. For example, to introduce the material cov- hands-on activities during classroom time and a greater
ered and capture student interest, each chapter begins reliance on student learning of core material outside of the
with an engaging story (a "Case Study," as described classroom. Ecology serves this purpose well with its clear,
 Preface IX

easy-to-read, and well-organized presentation of material. Case Studies Each chapter opens with an interesting vi-
In addition, there are several quantitative features-Ana- gnette-a Case Study. By presenting an engaging story or
lyzing Data exercises in the text and on the companion interesting application, the Case Study captures the read-
website, Hone Your Problem-Solving Skills and Hands- er 's attention while introducing the topic of the chapter.
On Problem Solving exercises-that can serve as the basis Later, the reader is brought full circle with the correspond-
for hands-on exercises. ing "Case Study Revisited" section at chapter's end. Each
Case Study relates to multiple levels of ecological hierar-
chy, thereby providing a nice lead-in to the Connections in
New to Ecology, Fourth Edition Nature feature, described next.
In striving to make Ecology the best teaching tool possi-
ble, we have updated, replaced, and edited sections of the Connections in Nature In most ecology textbooks, con-
text as appropriate. To accommodate the new features de- nections among levels of the ecological hierarchy are dis-
scribed below and keep the book to a manageable length, cussed briefly, perhaps only in the opening chapter. As a
we have revised and cut some sections. The Fourth Edition result, many opportunities are missed to highlight for stu-
includes: dents the fact that events in natural systems really are in-
terconnected. To facilitate the ability of students to grasp
Analyzing Data Exercises As part of their education how events in nature are interconnected, each chapter of
students should become comfortable working with and Ecology closes with a section that discusses how the mate-
analyzing data. Toward that end, we've added additional rial covered in that chapter affects and is affected by inter-
Analyzing Data exercises on the companion website. These actions at other levels of the ecological hierarchy. Where
online exercises give students extra practice with essential appropriate, these interconnections are also emphasized
skills such as performing calculations, making graphs, de- in the main body of the text.
signing experiments, and interpreting results.
Online Climate Change Connection Climate change has
Climate Change Connection Recognizing the increasing broad ecological effects with important implications for
evidence for and effects of climate change on ecological sys- conservation and ecosystem services. Half of the Fourth
tems, Climate Change Connection examples have been added Edition chapters include an online climate change exam-
into the main text of the book. These vignettes help students ple. These Climate Change Connection vignettes link top-
appreciate the many consequences of global climate change ics in the text to other levels of the ecological hierarchy,
on the distributions and functions of organisms as well as while enriching the student's understanding of ongoing
the ecosystems they depend on. Nearly half of the chapters climate change.
now include an in-text Climate Change Connection.
Ecological Inquiry Our understanding of ecology is con-
Hone Your Problem-Solving Skills The Fourth Edition stantly changing due to new observations and new results
features a new section added to the set of review ques- from ecological experiments and models. All chapters of
tions at the end of each chapter. The Hone Your Problem- the book emphasize the active, inquiry-based nature of
Solving Skills questions expose students to hypothetical what is known about ecology. This occurs throughout
situations or existing data sets, and allow them to work the narrative and is further highlighted by the Analyzing
through data analysis and interpretation to better under- Data exercises discussed earlier, and by the Figure Legend
stand key ecological concepts and relate these concepts Questions (described below). In addition, Ecology includes
to real life situations. a similar online Analyzing Data exercise for every in-text
Analyzing Data exercise, and hands-on interpretative and
quantitative exercises, described next.
Hallmark Features
In addition to the changes we just described, we've re- Hands-On Problem Solving Exercises This popular
vised and strengthened the key pedagogical features of feature of the Companion Website asks students to ma-
Ecology, introduced in previous editions: nipulate data, explore mathematical aspects of ecology
in more detail, interpret results from real experiments,
Pedagogical Excellence Students taking their first course and analyze simple model systems using simulations.
in ecology are exposed to a great deal of material, on a Each chapter of the book includes one or more Hands-
conceptual as well as individual-systems level. To help On Problems. These inquiry exercises can be used in two
them manage this vast amount of information, each chap- important ways: assigned as homework (all are avail-
ter of Ecology is organized around a small number of Key able to students via the Companion Website), or used
Concepts that provide up-to-date summaries of funda- as in-class exercises (each is provided in a new instruc-
mental ecological principles. tor format that makes it easy to incorporate them into
X Preface

classroom sessions as active learning exercises or discus- time as we respond to new discoveries and new ways of
sion topics) . teaching. We would love to hear from you-what you like
about the book, what you don't like, and any questions
Figure Legend Questions Each chapter includes 3--6 Fig- or suggestions you may have for how we can improve
ure Legend Questions that appear in maroon type at the the book. You can reach us individually or as a group
end of the legend. These questions encourage students to by sending an email message to [email protected], or
grapple with the figure and make sure they understand its by writing us at Ecology, Sinauer Associates, PO Box 407,
content. The questions range from those that test whether Sunderland, MA 01375 USA.
students understand the axes or other simple aspects of
the figure to those that ask students to develop or evalu-
ate hypotheses. Acknowledgments
We wish to express our appreciation to the people at
Ecological Applications In recent years, ecologists have Sinauer Associates, with whom we worked closely during
increasingly focused their attention on applied issues. the writing but especially during the book's production.
Similarly, many students taking introductory ecology Andy Sinauer supported the plan for the book from its
are very interested in applied aspects of ecology. Thus, inception. He enthusiastically participated in every phase
ecological applications (including conservation biology) along the way. Danna Lockwood and Kathaleen Emer-
receive great attention in this book. Discussions of applied son did a terrific job guiding the book through its many
topics are woven into each chapter, helping to capture and stages of production. Lou Doucette did a superb job of
retain student interest. copyediting our manuscript. Elizabeth Morales provided
the beautiful illustrations. David Mcintyre and Martha
Ecological Toolkits Nearly half of the chapters include Lorantos, our photo editors, always managed to find ex-
an Ecological Toolkit, a box inset in the chapters that de- quisite images that enhance the information in the figures.
scribes ecological "tools" such as experimental design, Joan Gemme facilitated several rounds of page design.
remote sensing, GIS, mark-recapture techniques, stable We love her final version that you see here, as well as her
isotope analysis, DNA fingerprinting, and the calculation elegant cover design. Dean Scudder masterminded the
of species-area curves. entire marketing effort and Marie Scavotto produced the
attractive brochure. Mark Belk (Brigham Young Univer-
Links to Evolution Evolution is a central unifying theme sity) and Amy Arnett (Unity College) expertly wrote and
of all biology, and its connections with ecology are very assembled the engaging online and instructor resources.
strong. Yet, ecology textbooks typically present evolution These resources, along with the impressive array of sup-
almost as a separate subject. As an alternative to the stan- plements listed on pp. xiv and xv, were coordinated by
dard approach, the first chapter of Ecology's Unit 2 (Chap- Jason Dirks.
ter 6) is devoted to describing the joint effects of ecology And finally, we'd like to thank some of the many peo-
and evolution. This chapter explores the ecology of evo- ple who helped us tum our ideas into a book in print. We
lution at both the population level and as documented are grateful to our colleagues who generously critiqued
in the sweeping history of life on Earth. Other topics in the plan for the book or read one or more chapters in man-
evolutionary ecology are explored in Chapter 7 (Life His- uscript; they are listed on the following pages. Among the
tory) and in Chapter 8 (Behavioral Ecology). Concepts or hundreds of people we contacted while researching this
applications that relate to evolution are also described in book, we also wish to thank the following individuals for
many other chapters. their special efforts in providing guidance and generously
sharing their time and expertise: Jocelyn Aycrigg, Jenifer
Art Program Many of Ecology's illustrations feature "bal- Hall-Bowman, John Jaenike, Michelle Koo, Karen Mabry,
loon captions," which tell a story that can be understood Debra VamVikites, and Tim Wright.
at a glance, without relying on the accompanying text.
WILLIAM D. BOWMAN
The art program is available as part of the Instructor 's
[email protected]
Resource Library (see Media and Supplements).
SALLY D. HACKER
[email protected]
Ecology Is a Work in Progress
MICHAEL L. CAIN
This book, like the subject we write about, does not con- [email protected]
sist of a set of unchanging ideas and fixed bits of infor-
MARCH2017
mation. Instead, the book will develop and change over
Reviewers

Reviewers for the Fourth Edition Ted Stankowich, California State University, Long Beach
Diana Tomback, University of Colorado, Denver
Anita Baines, University of Wisconsin, La Crosse
Monica Turner, University of Wisconsin, Madison
Nate Bickford, University of Nebraska at Kearney
Betsy Von Holle, University of Central Florida
Brian Butterfield, Freed-Hardeman University
Tobias Ziist, Cornell University
Kathleen Curran, Wesley College
John Fauth, University of Central Florida
Natalie Hyslop, University of North Georgia Reviewers for the First and Second Editions
Taegan McMahon, The University of Tampa David Ackerly, University of California, Berkeley
Luis Ruedas, Portland State University Gregory H. Adler, University of Wisconsin, Oshkosh
Kathleen Schnaars Uvino, University of Jamestown Stephana Allesina, University of Chicago
Stuart Allison, Knox College
Reviewers for the Third Edition Kama Almasi, University of Wisconsin, Stevens Point
Diane Angell, St. Olaf College Peter Alpert, University of Massachusetts, Amherst
David Armstrong, University of Colorado
Anita Baines, University of Wisconsin, La Crosse
Robert Baldwin, Clemson University
Sarah Dalrymple, University of California, Davis
Betsy Bancroft, Southern Utah University
Mark A. Davis, Macalester College
Jeb Barrett, Virginia Polytechnic Institute and
Abby Grace Drake, Skidmore College
State University
Joseph D'Silva, Norfolk State University
James Barron, Montana State University
Bret D. Elderd, Louisiana State University
Christopher Beck, Emory University
Mara Evans, University of California, Davis
Beatrix Beisner, University of Quebec at Montreal
Stephanie Fore, Truman State University
Mark C. Belk, Brigham Young University
Johanna Foster, Wartburg College
Michael A. Bell, Stony Brook University
Jennifer Fox, Georgetown University
Eric Berlow, University of California, Merced
Christiane Healey, University of Massachusetts, Amherst
Kim Bjorgo-Thorne, West Virginia Wesleyan College
Hopi Hoekstra, Harvard University
Charles Blem, Virginia Commonwealth University
John Jaenike, University of Rochester
Steve Blumenshine, California State University, Fresno
Piet Johnson, University of Colorado, Boulder
Carl Bock, University of Colorado
Melanie Jones, University of British Columbia
Daniel Bolnick, University of Texas, Austin
Gregg Klowden, University of Central Florida
Michael Booth, Principia College
Michelle Koo, University of California, Berkeley
April Bouton, Villanova University
Karen Mabry, New Mexico State University
Steve Brewer, University of Mississippi
A. Scott McNaught, Central Michigan University
David D. Briske, Texas A&M University
Rick Paradis, University of Vermont
Judie Bronstein, University of Arizona
Karin Pfennig, University of North Carolina, Chapel Hill
Linda Brooke Stabler, University of Central Oklahoma
Alysa Remsburg, Unity College
Kenneth Brown, Louisiana State University
Natalia Rybczynski, Canadian Museum of Nature
Rami Burks, Southwestern University
Dov Sax, Brown University
Stephen Burton, Grand Valley State University
Tom Schoener, University of California, Davis
Aram Calhoun, University of Maine
Andy Sih, University of California, Davis
XII Reviewers

Mary Anne Carletta, Georgetown College Vedham Karpakakunjaram, University of Maryland

Walter Carson, University of Pittsburgh Michael Kinnison, University of Maine
Peter Chabora, Queens College, CUNY Timothy Kittel, University of Colorado
David D. Chalcraft, East Carolina University Jeff Klahn, University of Iowa
Gary Chang, Gonzaga University Astrid Kodric-Brown, University of New Mexico
Colin A. Chapman, University of Florida Tom Langen, Clarkson University
Elsa Cleland, University of California, San Diego Jennifer Lau, Michigan State University
Cory Cleveland, University of Montana Jack R. Layne, Jr., Slippery Rock University
Liane Cochran-Stafira, Saint Xavier University Jeff Leips, University of Maryland, Baltimore County
Rob Colwell, University of Connecticut Stacey Lettini, Gwynedd-Mercy College
William Crampton, University of Central Florida Gary Ling, University of California, Riverside
James Cronin, Louisiana State University Scott Ling, University of Tasmania
Todd Crowl, Utah State University Dale Lockwood, Colorado State University
Anita Davelos Baines, University of Texas, Pan American Svata Louda, University of Nebraska
Andrew Derocher, University of Alberta Sheila Lyons-Sobaski, Albion College
Megan Dethier, University of Washington Richard Mack, Washington State University
John Ebersole, University of Massachusetts, Boston Lynn Mahaffy, University of Delaware
Erie Ellis, University of Maryland, Baltimore County Daniel Markewitz, University of Georgia
Sally Entrekin, University of Central Arkansas Michael Mazurkiewicz, University of Southern Maine
Jonathan Evans, University of the South Andrew McCall, Denison University
John Faaborg, University of Missouri Shannon McCauley, University of Michigan
William F. Fagan, University of Maryland Mark McPeek, Dartmouth College
Jennifer Fox, Georgetown University Scott Meiners, Eastern Illinois University
Kamal Gandhi, University of Georgia Bruce Menge, Oregon State University
Rick Gillis, University of Wisconsin, La Crosse Thomas E. Miller, Florida State University
Thomas J. Givnish, University of Wisconsin Sandra Mitchell, Western Wyoming College
Elise Granek, Portland State University Gary Mittelbach, Kellogg Biological Station,
Martha Groom, University of Washington Michigan State University
Jack Grubaugh, University of Memphis Russell Monson, University of Colorado
Vladislav Gulis, Coastal Carolina University Daniel Moon, University of North Florida
Jessica Gurevitch, Stony Brook University David Morgan, University of West Georgia
Bruce Haines, University of Georgia William F. Morris, Duke University
Nelson Hairston, Cornell University Kim Mouritsen, University of Aarhus
Jenifer Hall-Bowman, University of Colorado Shannon Murphy, George Washington University
Jason Hamilton, Ithaca College Courtney Murren, College of Charleston
Christopher Harley, University of British Columbia Shahid Naeem, Columbia University
Bradford Hawkins, University of California, Irvine Jason Neff, University of Colorado
Christiane Healey, University of Massachusetts, Amherst Scott Newbold, Colorado State University
Mike Heithaus, Florida International University Shawn Nordell, Saint Louis University
Kringen Henein, Carleton University, Ontario Timothy Nuttle, Indiana University of Pennsylvania
Kevin Higgins, University of South Carolina Mike Palmer, Oklahoma State University
Nat Holland, Rice University Kevin Pangle, The Ohio State University
Stephen Howard, Middle Tennessee State University Christopher Paradise, Davidson College
Randall Hughes, Florida State University Matthew Parris, University of Memphis
Vicki Jackson, Central Missouri State University William D. Pearson, University of Louisville
John Jaenike, University of Rochester Jan Pechenik, Tufts University
Bob Jefferies, University of Toronto Keith Pecor, The College of New Jersey
Art Johnson, Pennsylvania State University Karen Pfennig, University of North Carolina
Jerry Johnson, Brigham Young University Jeff Podos, University of Massachusetts, Amherst
Pieter Johnson, University of Colorado David M. Post, Yale University
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 Reviewers XIII

Joe Poston, Catawba College Jonathan Shurin, University of California, San Diego
Andrea Previtalli, Cary Institute of Ecosystem Studies Frederick Singer, Radford University
Seth R. Reice, University of North Carolina Richard Spellenberg, New Mexico State University
Alysa Remsburg, Unity College John J. Stachowicz, University of California, Davis
Heather Reynolds, Indiana University, Bloomington Christopher Steiner, Wayne State University
Jason Rohr, University of South Florida Cheryl Swift, Whittier College
Willem Roosenburg, Ohio University, Athens Ethan Temeles, Amherst College
Richard B. Root, Cornell University Michael Toliver, Eureka College
Scott Ruhren, University of Rhode Island Bill Tonn, University of Alberta
Nathan Sanders, University of Tennessee Kathleen Treseder, University of Pennsylvania
Mary Santelmann, Oregon State University Monica Turner, University of Wisconsin
Tom Sarro, Mount Saint Mary College Thomas Veblen, University of Colorado
Dov Sax, Brown University Don Waller, University of Wisconsin
Maynard H. Schaus, Virginia Wesleyan College Carol Wessman, University of Colorado
Sam Scheiner Jake F. Weltzin, University of Tennessee
Thomas Schoener, University of California, Davis Jon Witman, Brown University
Janet Schwengber, SUNY Delhi Stuart Wooley, California State University, Stanislaus
Erik P. Scully, Towson University Brenda Young, Daemen College
Catherine Searle, Oregon State University Richard Zimmerman, Old Dominion University
Dennis K. Shiozawa, Brigham Young University
Media and Supplements
to accompany Ecology, Fourth Edition

For the Student optimized for use in presentations. Complex figures

are provided in both whole and split versions.
Companion Website (ecology4e.sinauer.com)
• PowerPoint Resources: Three different PowerPoint
The Ecology Companion Website offers students a wealth
presentations are provided for each chapter of the
of study and review material. Access to the site is includ-
textbook:
ed with each new copy of the textbook (see inside front
cover). The Companion Website includes: • Figures: All figures and tables from each chapter,
with titles and full captions (in the Notes field) .
• Chapter Outlines and Summaries • Lecture: A complete lecture outline, including
• Hands-On Problem Solving Exercises that provide selected figures.
practical experience working with experimental • Hands-On Problems: Instructor versions of all
data and interpreting results from simulations and Hands-On Problems are provided in ready-to use
models. presentations.
• Analyzing Data Exercises----<ompanions to the • Hands-on Problem Solving Exercises: In addition
in-book exercises that give students extra practice to the instructor versions mentioned above, the stu-
with quantitative skills. dent versions of the exercises from the Companion
• Climate Change Connection examples explore Website, with answers, are included in Microsoft
the links between ecological concepts and climate Word format.
change. • Answers to Analyzing Data Exercises: Complete
• Web Extensions expand on the coverage of selected answers for all of the online Analyzing Data com-
topics introduced in the textbook. panion exercises.
• Online Quizzes are a great way for students to Test Bank
check their comprehension of the material covered The Ecology, Fourth Edition Test Bank includes a thorough
in each chapter (instructor registration required). set of questions for each chapter. All important concepts
• Web Stats Review-a brief statistics primer for are covered, and additional quantitative questions have
ecology. been added for the Fourth Edition. Each question is ref-
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 Media and Supplements XV

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Contents

CHAPTER 1 • The Web of Life 2 CONCEPT 1.3 Ecologists evaluate com peting hyp otheses
about natural system s w ith observations, experiments,
Deformity and Decline in Amphibian Populations: and models. 13
ACaseStudy 2 Answering Ecological Questions 13
Introduction 3 CLIMATE CHANGE CONNECTION
Approaches Used To Study Global Warming 14
CONCEPT 1.1 Events in the natural world are
interconnected . 3 • ECOLOGICAL TOOLKIT 1.1
Designing Ecological Experiments 15
Connections in Nature 3
ANALYZING DATA 1.1
CONCEPT 1.2 Ecology is the scientific study of Are Introduced Predators a Cause of Amphibian
interac tion s between organisms and their Decline? 17
en vironm ent. 8
What Is Ecology? 8 A CASE STUDY REVISITED Deformity and Decline in
Amphibian Populations 18
CONNECTIONS IN NATURE
Mission Impossible? 18

Unit 1
Organisms and Their Environment 21

CHAPTER 2 • The Physical Environment 22 Global Climate Patterns 31

Climate Variation and Salmon Abundance: CONCEPT 2.4 Region al climates reflect the
influen ce of ocean s and continents, m ountains, and
A Case Study 22
vegetation. 34
Introduction 23 Regional Climate Influences 34
CONCEPT 2.1 Climate is the most fundamental ANALYZING DATA 2.1
component of the physical environment. 23 How Do Changes in Vegetation Cover Influence
Climate 23 Climate? 37

CONCEPT 2.2 Winds and ocean currents result CONCEPT 2.5 Season al and long-term climate variation
from d ifferences in solar radiation across Earth's are associated w ith ch anges in Earth's position relative
su rface. 26 to the sun. 38
Atmospheric and Oceanic Circulation 26 Climate Variation over Time 38
CONCEPT 2.3 Large-scale atmosph eric and oceanic
circ ulation patterns establish global patterns of
temperature and precip itation. 31
 Contents XVII

CONCEPT Z.6 Salinity, acidity, and oxygen CONCEPT 4.Z The temperature of an organism is
concentrations are major determina nts of the chemical determined by exchanges of energy w ith the externa l
env ironment. 44 environment. 88
The Chemical Environment 44 Variation in Temperature 88
A CASE STUDY REVISITED Climate Variation and Salmon ANALYZING DATA 4.1
Abundance 46 How Does Fur Thickness Influence Metabolic Activity in
Endotherms? 97
CONNECTIONS IN NATURE
Climate Variation and Ecology 47 CONCEPT 4.3 The water balance of an organism is
determ ined by exchanges of water and solutes with the
externa l env ironment. 98
CHAPTER 3 • The Biosphere SO Variation in Water Availability 98
The American Serengeti-Twelve Centuries of A CASE STUDY REVISITED Frozen Frogs 105
Change in the Great Plains: A Case Study SO CONNECTIONS IN NATURE
Introduction 51 Desiccation Tolerance, Body Size, and Rarity 106
CONCEPT 3.1 Terrestrial biomes are cha racterized by the
growth forms of the dominant vegetation. 51 CHAPTERs • Coping with Environmental
Terrestrial Biomes 51 Variation: Energy 109
ECOLOGICAL TOOLKIT 3.1
Climate Diagrams 54 Toolmaking Crows: A Case Study 109
CLIMATE CHANGE CONNECTION Introduction 110
Tropical Forests and Greenhouse Gases 57
CONCEPT 5.1 Organ is ms obtain energy from sunlight,
ANALYZING DATA 3.1 from inorganic chemical compounds, or through the
How Will Climate Change Affect the Grasslands con sumption of organic compounds. 110
Biome? 6Z Sources of Energy 110
CONCEPT 3.Z Biological zones in freshwater ecosystems CONCEPT S.Z Radiant and chemical energy captured by
are associated w ith the velocity, depth, temperature, autotrophs is converted into stored energy in carbon-
clarity, and chemistry of the water. 71 carbon bonds. 112
Freshwater Biological Zones 71 Autotrophy 112
CONCEPT 3.3 Marine biological zones are determined by ANALYZING DATA 5.1
ocean depth, light availability, and the stab ility of the How Does Acclimatization Affect Plant Energy
bottom substrate. 73 Balance? 116
Marine Biological Zones 73 CONCEPT 5.3 Env ironmental constraints have resulted
A CASE STUDY REVISITED The American Serengeti- in the evolution of biochemical pathways that improve
'I'welve Centuries of Change in the Great Plains 79 the efficiency of photosynthesis. 117
RlJ CONNECTIONS IN NATURE Photosynthetic Pathways 117
~ Long-Term Ecological Research 80 CONCEPT 5.4 Heterotrophs have adaptations for
acquiring and assimilating energy efficiently from a
CHAPTER 4 • Coping with Environmental variety of organic sources. 123
Heterotrophy 123
Variation: Temperature and Water 84
• ECOLOGICAL TOOLKIT 5.1
Frozen Frogs: A Case Study 84 Stable Isotopes 1Z4
Introduction 85 A CASE STUDY REVISITED Toolmaking Crows 129
CONCEPT 4.1 Each species has a range of environmental CONNECTIONS IN NATURE
tolerances that de termines its potential geographic Tool Use: Adaptation or Learned Behavior? 130
distribution. 85
Responses to Environmental Variation 85
Exploring the Variety of Random
Documents with Different Content
 great efforts have been crowned with success, and
already the hour is at hand when Russia with her splendid
allies will finally crush by one joint and daring effort the last
resistance of the enemy.
A war such as this, unknown in history, must be continued
to the final and definite victory. Whoever dreams of peace
or desires it—at this moment—is a traitor to his country
and yields it to the enemy.
Carry out your duty, protect our beloved and glorious
country, submit yourselves to the Provisional Government,
render obedience to your chiefs, and remember that any
slackness in your service means a gain to your enemies.
With the firm conviction that the boundless love that you
have for our great country will ever remain in your hearts,
I pray that God may bless you, and that St. George the
Great Martyr may lead you to victory.
(Signed) Nicholas
(Countersigned) Alexeiev, C.G.S.
Father’s speech was a heavy blow to his men; they were stunned
from the shock. One of his personal guards had a heart attack while
Father was speaking. Father urged the troops to continue the war, at
any price, until victory was complete. The hour was at hand, he said,
when Russia would finally crush the last resistance of the enemy.
But this resistance did not materialize, nor, in fact, did the Emperor’s
own planned powerful attack. Some people of the upper classes in
Petrograd failed to support him in regard to this final blow against the
enemy, designed to win the war within three months, as Father had
promised in his last speech before the Duma in Petrograd a short
while before. It was not Father’s war, but the war of the nation. When
Uncle Michael came to urge Father to return immediately to G.H.Q.,
he found Father on the verge of departure, despite the serious
condition of Alexei.
The current malicious gossip was more important to the political
leaders than was the destiny of their country. These same leaders
afterwards criticized Father, saying that because he loved his family
so much, fear of having his family killed caused him to abdicate. The
killing of his family would not have solved the problem, which would
have been solved easily by the leaders themselves had they not
supported the traitors and not spread unfounded lies, especially
about my Father, whose men were shedding their blood at the front.
If I may ask, in what had we children sinned before our people that
we should have to give up our lives—we children who, from the
oldest to the youngest, denied ourselves all amusement and devoted
our energy to the war effort? Should we children have had to pay the
penalty of death? Why did not those busy conspirators take their
weapons and go to fight the common enemy?
They could not say anything against my older sisters, upon whom
they had never had an opportunity to lay their eyes. What they said
about me, because of my innocent jokes and pranks, I have never
given a rap about.
When Dr. Botkin read Father’s last addresses, his eyes moistened
and he added, “Only His Imperial Majesty could speak such deep
words.” Father repeated the words of Tsar Nicholas I: “Gde raz
podniat Russkii flag, on uzhe spuskatsia ne dolzhen.” (Where once
the Russian flag is raised, it shall never be lowered.)
Little by little we received more details. At first Father abdicated in
favor of Alexei, with Uncle Michael as Regent. But after consulting
Dr. Fedorov about his son’s delicate health, Father abdicated also for
Alexei in favor of his brother Michael, who was called in from
Gatchina to Petrograd, to accept first the Regency, and then the
Throne. To Father’s intense disappointment Uncle Misha, after
accepting, soon abdicated because of pressure by the Provisional
Government, in whose hands the nation’s fate then rested. This new
government placed under arrest many ministers and high officials,
because they refused to sign the loyalty pledge to the Provisional
Government. What was happening in Petrograd at this time Father
did not know until he reached home.
General Ruzsky and others, in order to hide their crime, spread the
most heartless rumor that Father had been drinking wine before his
abdication. However, Father was not alone during this painful ordeal.
With him on the train was his friend, the Minister of the Household of
the Court, Prince Vasily Dolgorukov; Flag Captain Nilov, A.D.C.;
General Voyeykov, Chief of Administration of the Palaces in
Tsarskoe Selo; Count Vladimir Fredericks, Father’s Chamberlain;
and the latter’s assistant, General Mossolov. Others present on the
train were A.D.C. Count Grabbe, Commandant of an Escort; A.D.C.
Captain Drenteln; A.D.C. Cyril Narishkin, head of the Chancery;
Colonel Mordvinov and General Dubensky. The Provisional
Government was represented by Kerensky’s friend, Vershchinin, and
others.
Prince Dolgorukov spoke to us of this merciless rumor, saying that
Father at the time drank only tea and paced back and forth in his
study. Father’s valet and his butler, after reading this lie, saw Count
Benckendorff and in tears said that those who spread such a lie had
committed a great sin toward their Emperor. It was also said that
during the last night on the train Father sat up all night in his study.
At midnight one of the engineer officers, who previously had
conducted many trips during Father’s travels, asked to see Father
and was received. He sank to his knees, kissed Father’s hand and
tearfully said, “Your Majesty, I will never serve these bandits. It is the
end of my life.” After the train reached the pavilion in Tsarskoe Selo
and just as Father was getting into his automobile and as the
standing officers were saluting Father for the last time, a shot was
heard and the engineer officer fell dead.
 PART III
Arrest And Exile
XIV
ARREST
It was decided to separate Mother from the rest of the family. But
Father objected, saying that it would be cruel to take Mother from her
sick children. At last it was conceded that Father should stay
downstairs in his apartment and Mother on the second floor with us.
Now that we all were under arrest, Mother was allowed to remain
with the sick children upstairs and Father was permitted to join his
family at mealtime. But all the conversation at the table had to be
carried on only in the Russian language. I was told that our private
wing of the palace was strongly guarded by a new kind of guard.
They were noisy and overbearing. Mother warned everybody to be
courteous to the sentry.
Had we children not been ill, Father would have insisted that we go
to England; at least the children if Mother had refused to leave. From
what he said later, we were convinced that he himself would not
have left Russia.
At once Kerensky appointed his communist friend Korovichenko,
whom he called his governor, as a commissar over us. This man
proved to be rude, dishonest, insulting, ignorant, and he quarrelled
with everyone in our household, disturbing and annoying us every
hour of the day. The guards were selected for the same qualities as
those of their masters. We were grateful when General Kornilov sent
Colonel Kobylinsky and Commissar Makarov to replace the cruel
Korovichenko.
From my sick bed all this was hard to imagine. I could not believe it,
until I was able to sit up by the window. I noticed that the appearance
and the action of the soldiers and officers were not the same.
Gradually our health improved. Tatiana had temporarily lost her
hearing and we had to write communications for her. Alexei’s
condition was still not up to par.
To kill our dull moments we played light games, worked on word
games and other puzzles and listened to French history read to us
by Mlle. Schneider.
We were told that Kerensky was coming to see us. The name
Kerensky brought terror to our hearts. Was he not the instigator
behind the treatment that Father was receiving? We waited,
dreading his visit. When the time of his arrival drew near, we were
filled with antagonism and fear. We were told that it was this enraged
man, Kerensky, and his communist friend, Korovichenko, who had
assembled our employees in the large hall. Kerensky told them that
they were no longer working for us, and that they were from now on
to take orders only from those who paid their wages. He meant of
course the Provisional Government, who appropriated the money
belonging to the people of Russia and us. But they themselves were
paid to live in an abundance such as one could only read about in
fiction.
Some of the servants were bribed to spy on us and Anna. A few
servants hated Anna to such an extent that they notified Kerensky of
Anna’s improved condition. Kerensky angrily ordered Anna to dress
at once, and she was then taken to prison. Alexei never before heard
such rough voices and he burst out crying. “Will he kill us?” he asked
his tutor. Those two men not only disturbed the family but upset
everybody. I saw these men passing by on their way to the
classroom where they were to see my parents. Father brought
Kerensky into a room and introduced my two older sisters, by merely
saying, “My daughters, Olga and Tatiana.” Marie and I were still in
bed, recovering from our illness.
When I first saw Kerensky, a man of medium height, he appeared to
be nervously twisting his finger. His face was pale and ugly, with
small greenish eyes imbedded in a peculiarly shaped head, which
was flat on top. His brown hair stood up similar to General Ruzsky’s.
Whenever I see a man like this, it always has an unpleasant
connotation for me. Outside in the hall and behind him, there was a
committee of workers, allegedly soldiers and sailors but really
nothing but released convicts, untidy, rough-looking individuals, who
were armed with daggers and hand grenades. We were frightened.
At first, I thought I was seeing this nightmare with my feverish eyes.
With the help of foreign spies, and convicts released by Kerensky
and the new leaders, these traitors were anxious to break the morale
of the armies. They told the soldiers that the land was to be divided
among them, but first they should return home in order to receive
their share of land. They said that it was a case of first come, first
served, and inasmuch as capital punishment had been abolished,
wholesale desertions from the armed forces resulted.
In the meantime the munitions and other factories fell into the hands
of the new government and many were set on fire. So the soldiers,
lacking supplies, had no choice but to desert their posts. All this
helped the enemy who was ready to collapse. Thanks to Kerensky
for the ruination of the army.
When the arrest of my family, especially of Father, became known to
the public, it caused a harmful effect on the morale of the Army, and
in general the situation became dangerous. Many soldiers left their
posts in panic; even in Tsarskoe Selo, Father’s own military escort,
not from choice but from fear, displayed red ribbons. The rebels were
freeing criminals who were breaking into wine cellars and becoming
intoxicated.
The snow was very heavy that winter, and the new leaders cared
little about having it cleared away. There was only a narrow space
available where Father and the others could walk about. This proved
actually to be a godsend to Father, because exercise was vital to his
health and was the only activity permitted him at this time. Each day
from the window I watched Father shovel snow. No doubt that for
Father it was much more than exercise. His physical exertions
enabled him to maintain his mental faculties in every respect.
In the beginning he had an unpleasant experience, during his
outdoor exercise, when he was allowed to walk in only a small area
close to the palace. One day on his return from his walk, he
extended his hand to one of the soldiers, but the man refused to take
it. This was hard on Father and made him realize the extent and
intensity of the propaganda against him. Father’s philosophical
attitude toward these incidents made him a greater hero in our eyes.
To us children, Kerensky was at first a beast, a dragon waiting to
devour us. His repeated visits kept us filled with terror. He thought at
first that he could come to the palace at any time and wander about
in our own home, without permission from the proper authority and
without Father’s consent. Kerensky, always accompanied by the
Marshal of the Court, followed by a messenger, was received in
Father’s study.
Father was always courteous, wishing to make everything easy for
the new government. But this man Kerensky at first did not seem to
know the meaning of courtesy. After a while he saw Father’s ready
cooperation and became quite human. We children began to feel
more relaxed in his presence. After several conversations with
Kerensky, our parents felt more confidence in him. However, Mother
could not forget the recent injustice she had suffered, and she hoped
her innocence was now proven. But Kerensky had made no attempt
to inform the public of the true situation at the palace. He was
responsible for Anna’s arrest, and Mother could not forgive him,
especially since she believed that Anna was still ill when she was
sent to prison. Madame Lili Dehn also had to leave the palace at the
same time. Moreover Madame Zizi Narishkina became ill with
pneumonia and left the palace because Mother felt she would
receive better care at a hospital.
This wise little lady was a favorite of my Father whom she had
known since his childhood. She was like a mother to our Mother. Her
kindness and simplicity was written all over her face. She insisted
that she be addressed as Madame instead of Princess, yet she was
a true-blooded Princess. Among us she called my Father Nicky. In
spite of her old age, she too had been a victim of unfair criticism. It
caused so much resentment that the newspapers had been forced to
retract their false stories.
Even my Aunt Olga, who loved peasants, had been condemned
because she enjoyed visiting and accepting their hospitality.
Our prison hours were well regulated. We were permitted two walks
during the day, between 11-12 and 2-5. With nervous excitement we
waited the designated hour, eager not to miss one second of the out-
of-doors. As the clock struck eleven, we and the staff gathered in the
semicircular room where we were to meet the guards who were to
accompany us on our walks. We had to wait sometimes as much as
one half hour. This meant our walk was sometimes curtailed that
much. We felt cheated, and the thought that Father could do nothing
about it made us feel worse. We discovered that the more we fretted,
the longer the delay; so we learned to wait submissively. The key to
the circular room was held by the commissar on duty, and the other
doors, including the balcony facing the Znamensky Cathedral and
the gate, were sealed. We had to wait until the commandant
appeared with the key to open the door. Since Korovichenko was as
a rule basking himself in the sun, he made it a habit of being late.
Even the sentries hated the sight of this man. When we did go out
finally, we walked briskly to cover lost ground. Sometimes we
crossed the bridge over the ravine, since the area was less exposed
to the public view. But because of demonstrations our afternoon
walks soon were scheduled later and we were outdoors until 8:00
P.M. Our friends helped us with our garden work. M. Gilliard proved
well-nigh indispensable.
At first we had no news from our relatives. But we were pleasantly
surprised when several letters came to Father and to Tatiana from
Aunt Xenia at Ai-Todor in the Crimea. The letters giving us news of
the family were the only joy we had had since our arrest. Our mail
was censored and parts of the letters we received were inked out.
Probably Aunt Xenia found our letters dry and uninteresting. But we
wrote them reluctantly, as in a daze—we so strongly felt the blow of
the happenings in our country and to its people. The shock was so
great that no outsider ever will know the feeling unless he lives
through a similar experience, that is, if he loved his country. We were
glad Granny was there too. It was her first stay in the Crimea since
the death of her husband, Alexander III. We had been wanting to
write to them but we feared repercussions.
For months we waited for a reply to our letters which we were
obliged to leave unsealed for censorship. We hoped that our coming
departure would take us to the Crimea to be together with the rest of
the family. Father hungrily read the newspapers, even though the
ones he received had been thoroughly censored, parts of them inked
out. All too often the papers were withheld altogether. Father read
every word carefully to find some clue to the real truth. One of our
best sources of information came from the various members of our
household. Before long they too no longer were free to leave the
palace. All around our home there was a strong guard posted,
especially in the small garden in front of Mother’s windows and the
garden gate across from the Znamensky Cathedral, which was the
closest to Mother’s balcony. She was very much disturbed, having
these men watch her windows; she never could look out. During this
time we saw some one drive in and out of the courtyard in Father’s
favorite automobile, a Packard sledge (sleigh). Kerensky of course
often used our private cars, chauffeurs, and even our valuable
horses.
After midnight another unpleasant incident occurred. Several
uncouth revolutionists, headed by a Pole named Mstislavsky, came
from Petrograd. They proceeded to show their thievish authority by
ordering the telephone and telegraph lines from the outside
disconnected. After a great deal of bickering and fighting with the
guards, who would not let them enter the grounds, Mstislavsky broke
down the gate with his heavy artillery trucks. It caused such a noise
that a large number of persons collected on the avenue and loud
voices were heard from the angry crowd. This noise awakened the
household. The bandits forced themselves towards and into the
palace, in spite of everything. They were armed and threatened the
officers on duty, “Shoot us, or we will shoot you.” Count Benckendorff
and Prince Dolgorukov came down and told Mstislavsky and his
companions that it was impossible for them to talk to the Emperor.
The intruders were asked to produce the required permit from
General Kornilov, commanding the troops in the area at this time.
The officers on duty then said, “You arrest us, or we will arrest you.”
Notwithstanding, the bandits went upstairs to the gallery room. There
they encountered Father who was walking toward them. Without a
word the bandits ran away in terror.
Kerensky was held responsible for this incident, having revealed to
Mstislavsky the anticipated departure of the Imperial family for
England via Murmansk.
Subsequently we heard that Mstislavsky ordered many innocent
families shot, people who were in his way; also he and his friends
seized from them anything to their liking.
We shortly learned that all escape routes were already blocked.
Korovichenko’s transfer lifted our hopes. His place was filled by
Colonel Eugene Kobylinsky, an officer of the Imperial Guard. We
were thankful to General Kornilov for sending this wonderful man,
who served my family faithfully in spite of many dangers to his own
life. He was sincerely anxious to alleviate our condition and was
determined to save us. But with many hundreds of men in our guard
he was helpless, as any action of his would have drawn suspicion
upon him. He went with us to Tobolsk and stayed there until four
days before our departure, when he became dangerously ill.
Consequently, we did not see this kind man again before we left for
Ekaterinburg. Another well-disposed man was Commissar Makarov,
a very intelligent and cultured person with a great deal of tact and
knowledge of several languages.
He had a prison record for having killed a policeman. For this, this
fine looking man paid fifteen years of his young life before he was
released. He accepted his punishment as having been deserved. His
imprisonment had not embittered him, and his gentle, kind manners
had not changed.
Many of the guards we had known from our childhood. Whenever
possible Colonel Kobylinsky tried to have these men accompany us
during our walks, as these guards could not tolerate an abusive
attitude of others toward us. Nevertheless, he did not dare to defend
us. By this time the Provisional Government had become convinced
that Father had no intention of making trouble for them. Father
eagerly followed the course of the war and grieved at the way it was
going. If only he could have now had a part in it, even as a private!
There were secret messages suggesting our escape to Germany.
Father answered, “No, we shall not escape like convicts.” One day
Father was walking, Alexei was sitting on the bench, with his dog Joy
at his feet, and we sisters were a short distance from Father, when a
large enclosed car rapidly drove in and two young men in it wanted
Father and Alexei to get into the car and escape with them. Father
was very upset by it. He said, “Go at once.” Soon after more new
guards appeared.
During this time we became tired of eating cabbages and carrots.
We longed for something different. Those days no green vegetables
came out of the greenhouses. Father saw the rich fields for labor
lying open to us, and he was willing to work. So, in the spring, Father
suggested a vegetable garden be planted in an open space where
some trees had been cut down. We were all eager for outdoor
exercise. Count Fredericks talked it over with Colonel Kobylinsky
who gave permission to go ahead. We were now full of enthusiasm,
with plenty of ideas of fresh food as our objective.
Father began to work, and even Mother, for the first time, cheerfully
was willing to leave the house in the afternoon. She was wheeled in
her chair into the garden. She sat under a tree near the brook, while
the guards paced back and forth on the bridge. Mother seldom
walked those days. We planted the seeds and watered the vegetable
beds from a barrel. As I worked, I thought of the words in the
beautiful Russian song which said:

“The Christ had a garden, where many roses bloomed,

He watered them thrice a day to make a wreath for himself.”

Because of the demonstrations, we worked late in the day, often till

8:00 P.M. In the evening Father read to us while we were sewing or
knitting. When the first green shoots appeared, we were thrilled with
the thought of salads within a month. The seedlings grew into bushy
plants. The blossoms became tiny beans. In another week or so we
would have our first harvest from sixty luxuriant beds in all. Spring,
which always seems to hold a special appeal for all Russians, was
beautiful but sad for us.
 XV
SUBJUGATION
When the first days of July with their white nights were
approaching, a time when night is much like day in those northern
regions, when twilight spreads a kind of magic transparency in the
distant sky and woods, we saw creeping figures with shining
bayonets emerge from the bushes. They were watching the windows
of the prisoners. I wish I did not love that great country with so much
promise, whose soul lies in debris now and of which I cannot speak
without the feeling of a heavy weight on my chest.
Before the leaves came out, we withdrew to an area where we
thought we could not be seen so easily. That led us to the
greenhouses. We found them dreadfully neglected. No one had
taken care of the plants. The gardeners had been discharged or put
to work in some other capacity. Now many rare and valuable bushes
surrounding the colonnades were cut down against everybody’s
objection. The orders, we were told, came from Kerensky. Tears
were in Father’s eyes to see such destruction.
We realized that the iron fence which protected us from the outside
was now our prison wall. The driveway was a source of fear. Even
the bushes and the trees of our beloved park secreted spies who
watched every move we made. Even though we were accustomed to
isolation from the world by a cordon of police and military protection,
being surrounded by unfriendly guards was indeed depressing.
During the turbulent days, even the swans cried mournfully every
morning because they knew we were in the palace, and they felt that
something was wrong that we did not speak to them and feed them.
Even these majestic birds must have known....
Our food now was ice-cold, more so than ever before. Our kitchens
were in a separate building and the food was wheeled through the
long tunnel in large carriers which had to be opened for inspection.
Therefore the food cooled off before it reached our apartments.
Sometimes we played on the “Children’s Island.” On several
occasions Alexei went out in a rowboat with his toy sailboats, but
was not allowed to enter his little four-room playhouse where he
used to play with his cousins and young cadets. It was locked up and
his rowboat taken away, making him very unhappy.
We enjoyed our cycling, Father and Alexei on bicycles and we four
girls on velocipedes. In the afternoon even Mother went into the
woods and sat in the shade with her tapestry work, copying the
original pattern of her Hepplewhite chair, while Father, his officer
friends and others were cutting down the dead trees. We sisters
helped to carry the smaller pieces and built tall stacks for the wood
to dry during the summer months.
We heard that our friend Captain Nilov, whom we called “the little
admiral”, once a commandant on the “Standard” and later at G.H.Q.
with Father, had been arrested on order of Kerensky and shot
without trial. This because he said while at G.H.Q. that he would kill
General Ruzsky. Fortunately for Ruzsky, Captain Nilov was denied
this pleasure. General Ruzsky also had a cordial dislike for Captain
Nilov because the latter knew that Ruzsky was a traitor.
We already by this time had begun to enjoy our fresh vegetables.
We lingered and feasted our eyes on the beauty of nature which until
now had been taken for granted. Alexei, not yet thirteen, in the early
summer delighted in shedding his boots and wading into the
sparkling water up to his knees. It did not take much to satisfy him
during these trying times. He wished nothing more from those
heartless men than to be free to enjoy God’s given creation. How
fortunate we human beings are, to see and feel all the loveliness and
enjoy it to the utmost! How cruel too when men deny this privilege to
their fellowmen!
Later, when we were in Tobolsk, Alexei recalled the Children’s Island
and wistfully expressed the hope that he might be able to return to it
and wade in the water again. He spoke of his playrooms, his small
cars, and then all of a sudden he seemed to realize that these
reminiscences of former places, dear to his heart, caused only pain,
and he never spoke of them again.
Many days we saw curious strangers and friends being driven from
the fence. The people, knowing the time of our walks, gathered
along the fence; some even climbed on top of carriages to get a
better view of us. Especially on Sundays, there were a great many
watching us through the railing.
Often we saw familiar, friendly faces in the crowd, but we were afraid
to recognize them. Once I thought I saw near the fence some of the
Tolstoys—Marie and Elizabeth, also Pasha and her brother. Mother
in her youth had met from time to time some of the Tolstoy relatives
who lived abroad. Another time we saw friends from Petrograd and
some nurses from the Tsarskoe Selo hospitals.
Near Easter my parents were informed that about eighty servants
and workers employed in the palace were to be discharged. My
family was perturbed, because some servants had families who
depended on their earnings, and many of them had been with my
family from the time of Father’s marriage. Before these people, so
close to us, left, both parents thanked them for their past services
and each one of them was presented with a gold or silver medal.
Orders came that Count Benckendorff and all ladies in waiting, as
well as Prince Dolgorukov, should leave us. However, they were
permitted to stay without compensation.
During Lent we were allowed to have services in our private chapel,
but Father Vassiliev had to eliminate Father’s name from the ritual.
With trembling voice he stopped in confusion when he came to the
part in which he was supposed to say, “Long Life for the Imperial
Family.” I am sure that in his mind he added the omission to his own
satisfaction. When Father Vassiliev became ill, after much
negotiating Count Fredericks, the Court Minister, received
permission to have Father Belyayev, a deacon and four singers
come to the palace during the Easter holidays.
Palm Sunday services had been held in the palace chapel on the
ground floor at the fourth entrance of the building. During the
services we were carefully watched. The guards were secreted
behind the draperies and the altar. Father Belyayev seeing all this
could not control his emotions; his tears fell freely down his
vestment.
During the Holy Week two services a day were the only refreshing
moments in our new lives. Mother stood behind a large screen made
in her favorite purplish-blue crystal glass, which Father had given her
previously. Behind the screen was a small, cushioned, kneeling
stand, on top of which rested a Psalter. On the wall to the right, were
several religious paintings, inherited from her Mother Alice and
several gifts from her Granny, Queen Victoria. The Psalter was
searched when it was brought in and again when taken away. In the
small room on Mother’s right adjoining the chapel some Bibles were
kept. Mother was very much annoyed when a guard stood behind
her throughout the service. “Even in this holy place,” she said, “one
is deprived of a moment of meditation.”
On the day of the Lord’s Crucifixion the revolutionists decided to bury
their own dead. With the red flags flying and a band blaring forth with
the Marseillaise and Chopin’s Funeral March, the procession
advanced through the avenues of lime trees and stopped opposite
the circular balcony from which we could see them marching. Among
the dead they paraded were bodies taken from the cemetery,
including those who died in the cellar, which they had set on fire
during a drinking spree. But their evil scheme came to an end on this
day, when angry, black clouds darkened the sky, when a terrific hail
and wind storm furiously broke whole branches from the trees and
pelted the metal roof of the palace with large hailstones. Candles
were lit. When daylight returned, the courtyard was flooded, and
there were large cakes of ice in the water. All was quiet now.
The same screeching cry and the detested Funeral March that
should be reserved only for the dead was now heard every day. It
became annoying even to the sentries. Often they whistled
sarcastically as soon as the demonstrators appeared. I even heard
this abhorrent March in my sleep.
Saturday night the staff, the servants and others, several hundred in
all, were present at the midnight service in our chapel, which lasted
till an early morning hour. The procession, headed by the priest who
carried an icon, went through the rooms with the lighted candles and
the message: “Khristos Voskrese (Christ is Risen)”, bringing the
hope that our dark lives might be brightened.
We thought it was a sad Easter, but a worse one was to follow. On
this Easter morning the staff, the chief of the guards, the
commandant, officers, ladies in waiting and a few others assembled
in Father’s library to break the blessed bread. In the afternoon those
on duty assembled in Mother’s room for congratulations.
This year our relatives, and high ranking military men, Ministers, and
the representatives of foreign countries no longer were permitted to
come to congratulate us on this Holy Day. Queen Olga of Greece,
the sister of the Grand Duke Constantine Constantinovich,
miraculously entered the grounds but her entry into the palace was
prevented. Her kind message and the Easter egg were delivered to
us by an officer. A great many others came but were turned back at
the gate.
There were several birthdays during our five months of imprisonment
at Tsarskoe Selo. First came my Father’s. On his birthday, services
were held in the chapel. The words “Long Life for the Tsar” were still
missing; a sad day. Then Mother’s, then came Tatiana’s, and on
June 5th, Russian calendar, I became sixteen years old—the year I
should have been officially presented to the Russian court. But there
was no debut for me and I did not care. The family did their best to
make an occasion of it. A service was held in the chapel and I
received congratulations from all around me. A year earlier a design
made by Fabergé for my lavalier had been approved by my family, to
be made of diamonds and pearls. I was to receive it on this day.
Instead, these sixteen diamonds and sixteen pearls, one for each
birthday and one for each name day, were sewn into my clothes
when we left for Siberia. Not long after my own birthday came
Marie’s, and then just before our departure from Tsarskoe Selo came
Alexei’s.
More about our imprisonment. First, there was some improvement in
Mother’s health. However, during the hot spell in July her heart
condition became worse. She was forced to lead a quiet life. There
were no more separations, no more hospital work and no attacks of
haemophilia for Alexei. We all fell into a routine. Lessons had been
entirely neglected since our illness. Some of our instructors came
from outside. They taught in the gymnasium and in other schools.
Now it was no longer permissible for them to resume their former
duties. Monsieur Gilliard spent most of his time with Alexei, and in
general was most helpful in reorganizing the household with which
the others were inexpert to cope.
We had books galore to choose from, and several pianos for our
use. We resumed piano lessons—now with Anastasia Hendrikova;
previously they were given by Mr. Konrad. Father began to teach us
history, geography and natural science; Mother, religion; Baroness
Buxhoeveden, English; Dr. Botkin, Russian; M. Gilliard, French; Mlle.
Schneider, mathematics. There was only one thing lacking—
inspiration.
From the very first Kerensky barged into Father’s rooms without
warning, much to the disgust of the Court Marshal who followed him
angrily. Kerensky asked Father whether he would go to Germany if
the Kaiser would extend an invitation. Father disappointed him by
answering, “I shall never set foot on German soil. I have already
previously rejected the invitation.” A car supposed to carry us over
the border crashed into the fence as it tried to drive through the gate.
Even if an opportunity had presented itself, dozens of strong chains
that bound us to our Mother country would not let us leave Russia.
There was an offer that we leave Russia by way of Murmansk, but
Kerensky betrayed it to the revolutionists, even though we would not
have accepted the offer.
Suddenly Kerensky wanted General Kornilov’s resignation but the
General refused to comply with his order. Kerensky then told Father
that Kornilov was a traitor. What was Kerensky? I wonder now if the
General was not in Kerensky’s way! Kerensky also was against
Captain Count Kotsebue, the Commandant of the Palace who
formerly was an Uhlan Guard officer. Kerensky forced him to give up
his post in favor of Kerensky’s communist friend, Colonel
Korovichenko.
Once after a walk when Father was about to enter the palace, one of
the new guards stepped in front of him and barred his entrance.
Alexei from the open door saw what happened and burst out crying.
Another time Father was walking with Prince Dolgorukov in the park
when one of the new officers followed close behind and stepped on
his heel. Father turned suddenly and hit the officer with his walking
cane so hard that the officer bent double. After that none of them
tried this incivility again.
Years later I spoke by chance to a former Russian officer who said
the Emperor should have “prayed less but worked more.” But this
officer’s wife at that time remarked to her husband, “What kind of
officer were you? When you became ill with appendicitis you carried
on like an infant! Is that bravery?” My Father not only prayed and
worked but possessed the bravery of a hero. Every day he spent ten
to fourteen hours at his desk. There was no other man that worked
and fought harder and with more determination than did Father.
Such words were spoken only by German traitors and weaklings
who did everything to buy the privilege to stay far behind the front
lines. Some said that Father was mild. Perhaps he was; he might
have been appreciated in another country. However, many said that
Father should have ordered guns, but the Emperor would not take
the lives of men. He was very kind because he did not believe in the
ruthlessness of Ivan the Terrible, or Lenin, Trotsky, Apfelbaum,
Rosenfeld, Himmer, and others.
The very things we loved most, were now turning against us. Each
morning when I awakened I hoped for an improvement. But one
glance around convinced me the times were not right. One day a
bullet hit a window in Father’s study and left an ugly round scar,
showing the great thickness of the glass. Our walks continued, but
every time we went out we discovered many familiar faces had
disappeared, and were replaced by new ones. In the park the
sentries followed us closely and engaged us in conversation. Most
often we did not care to hear what they said. Then came an order
limiting our walk to the first bridge of the brook. Now we were
confined to a more restricted area. We tried to ignore the
impertinence of guards who lolled in our chairs on the lawn in front of
the small balcony which led into the entry room. This balcony was a
few steps up and had an entrance on each side, but the arbitrary
guards prevented us from entering except through one side only.
Mother watched us from the window, and when we returned to her,
one glance indicated that she had been weeping. We knew it was
the sight of our being so restricted that made her cry. Whether long
or short, these walks remained the most coveted events of our day.
Mother too was wheeled out in her chair, and sat with her
embroidery in the shade. Usually she was surrounded by the young
guards who asked her all kinds of questions, mostly religious ones.
Those big children understood her and she won them with her
kindness.
At first our captivity at Tsarskoe Selo was not so difficult, even
though there was a complete lack of privacy and our freedom to
come and go was restricted. In fact it was not much different from
our usual routine as we had been accustomed to watchful eyes and
many limitations. We would have been fairly comfortable if we had
had enough heat and the right food for the convalescents.
The familiar rooms, furnished with what we always considered to be
our own possessions, were at once comforting and disturbing. But
soon we found out that these things and many personal household
treasures no longer belonged to us. Many of these were confiscated
immediately, including all Mother’s silver sets. Some of these were
heirlooms from her Granny, Queen Victoria, and some were Father’s
wedding gifts. Other confiscations included trays, platters, urns and
numerous gold plates, gold tea glass holders and spoons, over five
hundred table place settings, Mother’s imperial jade figurines and
crosses by Bolin, the most famous jeweller in Petrograd; also many
priceless gold icons decorated with precious stones and other
treasures.
They even took Alexei’s jade and rock crystal collection of animals,
gold swords, miniatures of the family, and his icons, many of which
were presented to him during his illness in Spala in 1912 and were
especially esteemed by him, because the people had prayed before
them to spare his life. Some of these were in gold, studded in
precious stones. On his birth in 1904 the Shah of Persia ordered a