Ecology Basics

Ecology Basics
The Editors of Salem Press
SALEM PRESS
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MAGILL’S C H O I C E
Ecology Basics
Volume 1
Acid deposition—Lichens
Volume 2
Mammalian social systems—Zoos

Appendices
Indexes
edited by
The Editors of Salem Press
Salem Press, Inc.

Pasadena, California Hackensack, New Jersey
Copyright © 2004, by Salem Press, Inc.
All rights in this book are reserved. No part of this work may be used or
reproduced in any manner whatsoever or transmitted in any form or by
any means, electronic or mechanical, including photocopy, recording, or
any information storage and retrieval system, without written permission
from the copyright owner except in the case of brief quotations embodied
in critical articles and reviews. For information address the publisher, Sa-
lem Press, Inc., P.O. Box 50062, Pasadena, California 91115.
∞ The paper used in these volumes conforms to the American Na-

tional Standard for Permanence of Paper for Printed Library Materials,
Z39.48-1992 (R1997).
Library of Congress Cataloging-in-Publication Data

Ecology basics / edited by the editors of Salem Press.
p. cm. — (Magill’s choice)
Includes bibliographical references.
ISBN 1-58765-174-2 (set : alk. paper) — ISBN 1-58765-175-0 (v. 1 : alk.
paper) — ISBN 1-58765-176-9 (v. 2 : alk. paper)
1. Ecology—Encyclopedias. I. Salem Press. II. Series.
QH540.4.E39 2003
577′.03—dc21
2003011370
First Printing
printed in the united states of america

Contents of Volume 1
Publisher’s Note . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ix
Contributors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xiii
Complete List of Contents . . . . . . . . . . . . . . . . . . . . . . . . . xvii
Acid deposition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Adaptations and their mechanisms . . . . . . . . . . . . . . . . . . . . . . 7
Adaptive radiation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Allelopathy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Altruism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Animal-plant interactions . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Balance of nature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
Biodiversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
Biogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Biological invasions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Bioluminescence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
Biomagnification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Biomass related to energy . . . . . . . . . . . . . . . . . . . . . . . . . . 50
Biomes: determinants . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Biomes: types . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Biopesticides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
Biosphere concept . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Camouflage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
Chaparral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
Clines, hybrid zones, and introgression . . . . . . . . . . . . . . . . . . 80
Coevolution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
Colonization of the land . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
Communication . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
Communities: ecosystem interactions . . . . . . . . . . . . . . . . . . . 100
Communities: structure . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
Competition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111
Conservation biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
Convergence and divergence . . . . . . . . . . . . . . . . . . . . . . . . 120
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Ecology Basics
Deep ecology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123

Defense mechanisms . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125
Deforestation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
Demographics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
Dendrochronology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
Desertification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
Deserts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154
Development and ecological strategies . . . . . . . . . . . . . . . . . . 161
Displays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167
Ecology: definition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171

Ecology: history . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179
Ecosystems: definition and history . . . . . . . . . . . . . . . . . . . . 184
Ecosystems: studies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191
Endangered animal species . . . . . . . . . . . . . . . . . . . . . . . . . 196
Endangered plant species . . . . . . . . . . . . . . . . . . . . . . . . . . 205
Erosion and erosion control . . . . . . . . . . . . . . . . . . . . . . . . . 211
Ethology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
Eutrophication . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222
Evolution: definition and theories . . . . . . . . . . . . . . . . . . . . . 227
Evolution: history . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 236
Evolution of plants and climates . . . . . . . . . . . . . . . . . . . . . . 241
Extinctions and evolutionary explosions . . . . . . . . . . . . . . . . . 246
Food chains and webs . . . . . . . . . . . . . . . . . . . . . . . . . . . . 255

Forest fires . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 258
Forest management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 263
Forests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 269
Gene flow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 274

Genetic diversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278
Genetic drift . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281
Genetically modified foods . . . . . . . . . . . . . . . . . . . . . . . . . 284
Geochemical cycles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 288
Global warming . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 292
Grasslands and prairies . . . . . . . . . . . . . . . . . . . . . . . . . . . 298
Grazing and overgrazing . . . . . . . . . . . . . . . . . . . . . . . . . . 304
Greenhouse effect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 308
Habitats and biomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313

Habituation and sensitization . . . . . . . . . . . . . . . . . . . . . . . 319
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Contents
Herbivores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 326
Hierarchies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 329
Human population growth . . . . . . . . . . . . . . . . . . . . . . . . . 333
Hydrologic cycle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 338
Insect societies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 343

Integrated pest management . . . . . . . . . . . . . . . . . . . . . . . . 351
Invasive plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 354
Isolating mechanisms . . . . . . . . . . . . . . . . . . . . . . . . . . . . 358
Lakes and limnology . . . . . . . . . . . . . . . . . . . . . . . . . . . . 364

Landscape ecology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 374
Lichens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381
vii
Contents of Volume 2
Mammalian social systems . . . . . . . . . . . . . . . . . . . . . . . . . 385

Marine biomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 391
Mediterranean scrub . . . . . . . . . . . . . . . . . . . . . . . . . . . . 399
Metabolites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 402
Migration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 407
Mimicry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 415
Mountain ecosystems . . . . . . . . . . . . . . . . . . . . . . . . . . . . 419
Multiple-use approach . . . . . . . . . . . . . . . . . . . . . . . . . . . 422
Mycorrhizae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 425
Natural selection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 428

Nonrandom mating, genetic drift, and mutation . . . . . . . . . . . . . 435
Nutrient cycles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 440
Ocean pollution and oil spills . . . . . . . . . . . . . . . . . . . . . . . . 444

Old-growth forests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 452
Omnivores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 455
Ozone depletion and ozone holes . . . . . . . . . . . . . . . . . . . . . 457
Paleoecology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 464
Pesticides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 470
Pheromones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 476
Phytoplankton . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 482
Poisonous animals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 486
Poisonous plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 490
Pollination . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 495
Pollution effects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 500
Population analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 507
Population fluctuations . . . . . . . . . . . . . . . . . . . . . . . . . . . 513
Population genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 520
Population growth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 528
Predation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 536
Punctuated equilibrium vs. gradualism . . . . . . . . . . . . . . . . . . 543
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Ecology Basics
Rain forests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 549

Rain forests and the atmosphere . . . . . . . . . . . . . . . . . . . . . . 554
Rangeland . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 560
Reefs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 564
Reforestation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 572
Reproductive strategies . . . . . . . . . . . . . . . . . . . . . . . . . . . 576
Restoration ecology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 583
Savannas and deciduous tropical forests . . . . . . . . . . . . . . . . . 586

Slash-and-burn agriculture . . . . . . . . . . . . . . . . . . . . . . . . . 590
Soil . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 594
Soil contamination . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 601
Speciation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 604
Species loss . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 608
Succession . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 612
Sustainable development . . . . . . . . . . . . . . . . . . . . . . . . . . 618
Symbiosis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 621
Taiga . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 629
Territoriality and aggression . . . . . . . . . . . . . . . . . . . . . . . . 633
Trophic levels and ecological niches . . . . . . . . . . . . . . . . . . . . 641
Tropisms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 650
Tundra and high-altitude biomes . . . . . . . . . . . . . . . . . . . . . 655
Urban and suburban wildlife . . . . . . . . . . . . . . . . . . . . . . . . 659
Waste management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 667

Wetlands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 672
Wildlife management . . . . . . . . . . . . . . . . . . . . . . . . . . . . 677
Zoos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 681
Appendices
Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 687
Web Sites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 729
Indexes
Categorized Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 735
Subject Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 741
xxviii
Publisher’s Note
M agill’s Choice: Ecology Basics offers 132 essays, each of which covers a
fundamental ecological concept taught in biology, environmental
science, and introductory ecology courses. Alphabetically arranged, these
topics range from the level of individual organisms and their interactions
with the environment through populations of organisms and communities of
more than one species, to the level of ecosystems and global ecology. These
two volumes provide coverage of ecology in its broadest scientific sense:
not only according to Ernst Haeckel’s original definition—the interaction
of organisms with their environment—but also in the now-conventional
sense first expounded in 1954 by Herbert George Andrewartha and Louis
Charles Birch as the processes that influence the “abundance and distribu-
tion” of organisms, and, further, in the sense of ecosystem ecology, intro-
duced in 1971 by Eugene Odum. All levels taught in most general and in-
troductory courses, as well as key environmental issues and ecological
impacts of pollution, are considered here.
Previously appearing in three Salem publications—Magill’s Encyclope-
dia of Science: Animal Life (2002), Magill’s Encyclopedia of Science: Plant Life
(2003), and Earth Science (2001)—the essays begin by listing the subdis-
cipline of ecology into which the topic is generally categorized; where
the topic is central to more than one subdiscipline, all are listed. There
follows a brief synopsis defining the topic and its significance. The text
of each essay, ranging from two to six pages, is subheaded to flag the
core concepts addressed. Each essay ends with the signature of the acade-
mician who wrote it, a full set of cross-references to other essays in this
publication that treat related concepts, and a list of sources for further
study to assist students and general readers searching for fuller infor-
mation.
The subdisciplines into which the essays are classified are as follows:
• Agricultural ecology: Also called “agroecology,” the study of agri-
cultural ecosystems, their components (such as crop species), func-
tions, interactions, and impact on natural ecosystems and abiotic fac-
tors such as atmospheric and water systems—often with an emphasis
on the development of sustainable systems.
• Aquatic and marine ecology: The study of the ecology of freshwater
systems (rivers, lakes), estuaries, and marine environments (both
coastal and open ocean), including the physical, chemical, and bio-
logical processes associated with them.
ix
Ecology Basics
• Behavioral ecology: The study of how individual organisms interact

through behavior with other organisms and their environment to sur-
vive and reproduce, which has an impact on population.
• Biomes: The primary, large-scale ecosystems of the world, largely
identified with geographical regions and classified on the basis of
precipitation, temperature, climate, soil types, flora, and fauna.
• Chemical ecology: Concerns the biochemicals (or semiochemicals)
that organisms produce and release, which have physiological and
behavioral effects on other organisms.
• Community ecology: The study of the impacts that populations of
different species have on populations of other species with which
they interact, be those interactions of plants with other plants, ani-
mals with other animals, or plants with animals. The emphasis is on
how these populations of different species change, enhance, or de-
limit one another. Population ecology is related but is focused on
growth and change within populations of a single species.
• Ecoenergetics: The flow of energy through ecological systems at all
levels, from individual organisms, populations, and communities
to ecosystems and the global environment. Includes abiotic factors
(such as geochemical cycles) as well as biotic factors.
• Ecosystem ecology: The study of the flow of energy into, through,
and out of large-scale systems, and how that flow influences all
abiotic factors and living organisms in the ecosystem.
• Ecotoxicology: The study of natural and human-made pollutants and
their toxic effects on organisms, populations, communities, and eco-
systems, as well as the ways these pollutants impact ecological pro-
cesses to change ecosystems and their components.
• Evolutionary ecology: The study of how evolutionary processes such
as selection and adaptation influence the interactions of organisms
with their environments and shape species and ecosystems.
• Global ecology: The study of the impacts of such factors as global
warming, pollution, and disease on organisms and ecosystems world-
wide. Much of global ecology considers the ecological impacts of
human-driven influences such as international travel, trade, the built
environment, and the use of petrochemicals.
• History of ecology: The development of the discipline of ecology.
• Landscape ecology: The science of managing the habitat components
of modified landscapes—a burgeoning field concerned with preserv-
ing the naturalness of modified landscapes while minimizing the
negative impact of human intrusion in natural habitats within these
landscapes.
x
Publisher’s Note
• Paleoecology: The study of past ecosystems and environments.

• Physiological ecology: Sometimes called “autoecology,” “ecophysi-
ology,” or “comparative physiology,” a type of ecology that focuses
on individual organisms, examining how they function mechanically
and physiologically in their environments and how such factors as
temperature, seasons, soil, and nutrients affect survival and repro-
duction of those organisms. Unlike morphology or physiology, the
emphasis is on linking individual organisms, via their performance
attributes, to populations and communities.
• Population ecology: The study of the growth and decline of groups
of individuals of the same species, and how these fluctuations func-
tion in relation to other populations in the same ecosystem. Exam-
ines such factors as the availability of food and hence predation,
herbivory, and mutualisms. Community ecology is closely related to
population ecology but focuses on the interactions between popula-
tions of different species.
• Restoration and conservation ecology: Restoration ecology is the
study and implementation of ways to return degraded or deteriorat-
ing communities and ecosystems to their original condition. Restora-
tion ecologists work to restore habitat and return endangered species
to viable numbers; they do not seek to restore extinct species or re-
create ancient habitats. Conservation ecology is the use of biological
science to design and implement methods to ensure the survival of
species, ecosystems, and ecological processes. Conservation biolo-
gists develop strategies to preserve biodiversity before it becomes de-
graded.
• Soil ecology: The study of soil as an ecosystem, including the interac-
tions of both abiotic and biotic components of soil: water, minerals,
bacteria, fungi, plant matter, microbial organisms, and small animals
such as insects and worms. Soil ecology extends beyond the physical
borders of soil to include the impact of soil on aboveground life-
forms such as larger plants and animals, as well as processes (geo-
chemical cycles, erosion, human agricultural practices) that impact
soil.
• Speciation: The study of the processes whereby new species arise.
• Theoretical ecology: The study of the fundamental theories, con-
cepts, and models of ecological relationships, from the simplest pred-
ator-prey, host-parasite models to population, community, ecosys-
tem, global, and evolutionary models.
For convenience, both volumes of Ecology Basics contain a full list of the
contents. At the end of volume 2, several research tools are offered: a Glos-
xi
Ecology Basics
sary, a list of Web Sites, a Categorized Index (by type of ecology), and a
Subject Index.
All essays were prepared by qualified academicians and experts, with-
out whose invaluable contributions these volumes would not be possible.
Their names and affiliations follow.
xii
Contributors
Richard Adler David L. Chesemore
University of Michigan, Dearborn California State University, Fresno
Steve K. Alexander Sneed B. Collard

University of Mary Hardin-Baylor University of West Florida
Richard W. Arnseth J. A. Cooper

Science Applications International Independent Scholar
George K. Attwood Alan D. Copsey

Maharishi International University Central University of Iowa
David Landis Barnhill Mark S. Coyne

Guilford College University of Kentucky
Erika L. Barthelmess Greg Cronin

St. Lawrence University University of Colorado at Denver
Margaret F. Boorstein James F. Crow

C. W. Post College of Long Island University of Wisconsin
University
Gordon Neal Diem
P. E. Bostick ADVANCE Education and
Kennesaw State College Development Institute
Catherine M. Bristow John P. DiVincenzo

Michigan State University Middle Tennessee State University
William R. Bromer Allan P. Drew

Pepperdine University SUNY, College of Environmental
Science and Forestry
Steven D. Carey
University of Mobile Frank N. Egerton
University of Wisconsin, Parkside
Richard W. Cheney, Jr.
Christopher Newport University David K. Elliott
Northern Arizona University
xiii
Ecology Basics
Jessica O. Ellison David Wason Hollar, Jr.

Clarkson University Rockingham Community College
Danilo D. Fernando Robert Hordon

SUNY, College of Environmental Rutgers University
Science and Forestry
Richard D. Howard
James F. Fowler Independent Scholar
State Fair Community College
Jason A. Hubbart
Roberto Garza California State University, Fresno
San Antonio College
Samuel F. Huffman
Ray P. Gerber University of Wisconsin, River Falls
Saint Joseph’s College
Lawrence E. Hurd
D. R. Gossett Washington and Lee University
Louisiana State University,
Shreveport Diane White Husic
East Stroudsburg University
Jerry E. Green
Miami University Jeffrey A. Joens
Florida International University
Linda Hart
University of Wisconsin, Madison Christopher Keating
Angelo State University
Thomas E. Hemmerly
Middle Tennessee State University Kenneth M. Klemow
Wilkes University
John S. Heywood
Southwest Missouri State University P. R. Lannert
Independent Scholar
Joseph W. Hinton
Independent Scholar David M. Lawrence
John Tyler Community College
Carl W. Hoagstrom
Ohio Northern University Walter Lener
Nassau Community College
Virginia L. Hodges
Northeast State Technical W. David Liddell
Community College Utah State University
xiv
Contributors
Robert Lovely Edward N. Nelson

University of Wisconsin, Madison Oral Roberts University
Yiqi Luo Bryan Ness

University of Oklahoma Pacific Union College
Michael L. McKinney John G. New

University of Tennessee, Knoxville Loyola University of Chicago
Kristie Macrakis Edward B. Nuhfer

Harvard University University of Wisconsin, Platteville
Paul Madden Oghenekome U. Onokpise

Hardin-Simmons University Florida A&M University
Nancy Farm Männikkö Robert W. Paul

Independent Scholar St. Mary’s College of Maryland
Linda Mealey Rex D. Pieper

College of St. Benedict New Mexico State University
John S. Mecham Noreen D. Poor

Texas Tech University University of South Florida
Randall L. Milstein Robert Powell

Oregon State University Avila College
Eli C. Minkoff Donald R. Prothero

Bates College Occidental College
Richard F. Modlin Carol S. Radford

University of Alabama, Huntsville Maryville University, St. Louis
Thomas C. Moon P. S. Ramsey

California University of Pennsylvania Independent Scholar
Randy Moore C. Mervyn Rasmussen

Wright State University Independent Scholar
Christina J. Moose
Independent Scholar
xv
Ecology Basics
Ronald J. Raven Sanford S. Singer

State University of New York at University of Dayton
Buffalo
Elizabeth Slocum
Darrell L. Ray Independent Scholar
University of Tennessee, Martin
Dwight G. Smith
David D. Reed Southern Connecticut State University
Michigan Technological University
Roger Smith
Gregory J. Retallack Independent Scholar
University of Oregon
Valerie M. Sponsel
Mariana Louise Rhoades University of Texas, San Antonio
St. John Fisher College
Joan C. Stevenson
James L. Robinson Western Washington University
University of Illinois at Urbana-
Champaign Dion Stewart
Adams State College
David W. Rudge
Western Michigan University Toby R. Stewart
Independent Scholar
James L. Sadd
Occidental College Marshall D. Sundberg
Emporia State University
Lisa M. Sardinia
Pacific University Frederick M. Surowiec
Independent Scholar
Samuel M. Scheiner
Northern Illinois University Leslie V. Tischauser
Prairie State College
John Richard Schrock
Emporia State University Yujia Weng
Northwest Plant Breeding Company
Donna Janet Schroeder
College of St. Scholastica Samuel I. Zeveloff
Weber State College
Jon P. Shoemaker
University of Kentucky Ming Y. Zheng
Gordon College
xvi
Complete List of
Contents
Volume 1
Acid deposition, 1 Deforestation, 131

Adaptations and their Demographics, 137
mechanisms, 7 Dendrochronology, 145
Adaptive radiation, 12 Desertification, 149
Allelopathy, 15 Deserts, 154
Altruism, 18 Development and ecological
Animal-plant interactions, 24 strategies, 161
Balance of nature, 28 Displays, 167
Biodiversity, 32 Ecology: definition, 171
Biogeography, 37 Ecology: history, 179
Biological invasions, 40 Ecosystems: definition and
Bioluminescence, 43 history, 184
Biomagnification, 47 Ecosystems: studies, 191
Biomass related to energy, 50 Endangered animal species, 196
Biomes: determinants, 55 Endangered plant species, 205
Biomes: types, 59 Erosion and erosion control, 211
Biopesticides, 65 Ethology, 215
Biosphere concept, 69 Eutrophication, 222
Camouflage, 72 Evolution: definition and theories,
Chaparral, 76 227
Clines, hybrid zones, and Evolution: history, 236
introgression, 80 Evolution of plants and climates,
Coevolution, 86 241
Colonization of the land, 90 Extinctions and evolutionary
Communication, 95 explosions, 246
Communities: ecosystem Food chains and webs, 255
interactions, 100 Forest fires, 258
Communities: structure, 104 Forest management, 263
Competition, 111 Forests, 269
Conservation biology, 119 Gene flow, 274
Convergence and divergence, 120 Genetic diversity, 278
Deep ecology, 123 Genetic drift, 281
Defense mechanisms, 125 Genetically modified foods, 284
xvii
Ecology Basics
Geochemical cycles, 288 Human population growth, 333

Global warming, 292 Hydrologic cycle, 338
Grasslands and prairies, 298 Insect societies, 343
Grazing and overgrazing, 304 Integrated pest management, 351
Greenhouse effect, 308 Invasive plants, 354
Habitats and biomes, 313 Isolating mechanisms, 358
Habituation and sensitization, 319 Lakes and limnology, 364
Herbivores, 326 Landscape ecology, 374
Hierarchies, 329 Lichens, 381
Volume 2
Mammalian social systems, 385 Population analysis, 507

Marine biomes, 391 Population fluctuations, 513
Mediterranean scrub, 399 Population genetics, 520
Metabolites, 402 Population growth, 528
Migration, 407 Predation, 536
Mimicry, 415 Punctuated equilibrium
Mountain ecosystems, 419 vs. gradualism, 543
Multiple-use approach, 422 Rain forests, 549
Mycorrhizae, 425 Rain forests and the atmosphere,
Natural selection, 428 554
Nonrandom mating, genetic drift, Rangeland, 560
and mutation, 435 Reefs, 564
Nutrient cycles, 440 Reforestation, 572
Ocean pollution and Reproductive strategies, 576
oil spills, 444 Restoration ecology, 583
Old-growth forests, 452 Savannas and deciduous tropical
Omnivores, 455 forests, 586
Ozone depletion and Slash-and-burn agriculture, 590
ozone holes, 457 Soil, 594
Paleoecology, 464 Soil contamination, 601
Pesticides, 470 Speciation, 604
Pheromones, 476 Species loss, 608
Phytoplankton, 482 Succession, 612
Poisonous animals, 486 Sustainable development, 618
Poisonous plants, 490 Symbiosis, 621
Pollination, 495 Taiga, 629
Pollution effects, 500 Territoriality and aggression, 633
xviii
Complete List of Contents
Trophic levels and ecological Wildlife management, 677

niches, 641 Zoos, 681
Tropisms, 650
Tundra and high-altitude biomes, Glossary, 687
655 Web Sites, 729
Urban and suburban wildlife, 659
Waste management, 667 Categorized Index, 735
Wetlands, 672 Subject Index, 741
xix
ACID DEPOSITION
Types of ecology: Aquatic and marine ecology; Ecotoxicology
Electric utilities, industries, and automobiles emit sulfur dioxide and nitrogen ox-
ides that are readily oxidized into sulfuric and nitric acids in the atmosphere.
Long-range transport and dispersion of these air pollutants produce regional acid
deposition. Acid deposition alters aquatic—and possibly forest—ecosystems and
accelerates corrosion of buildings, monuments, and statuary.
I n 1872 Robert Angus Smith used the term “acid rain” in his book Air and
Rain: The Beginnings of a General Climatology to describe precipitation af-
fected by coal-burning industries. Today, “acid rain” refers to the deposi-
tion of acidic gases, particles, and precipitation (rain, fog, dew, snow, or
sleet) on the surface of the earth. The normal acidity of rain is pH 5.6, which
is caused by the formation of carbonic acid from water-dissolved carbon
dioxide. The acidity of precipitation collected at monitoring stations
around the world varies from pH 3.8 to 6.3 (pH 3.8 is three hundred times
as acidic as pH 6.3). The acidity is created when sulfur dioxide and nitro-
gen oxides react with water and oxidants in the atmosphere to form water-
soluble sulfuric and nitric acids. Ammonia, as well as soil constituents
such as calcium and magnesium that are often present in suspended dust,
neutralizes atmospheric acids, which helps explain the geographical varia-
tion of precipitation acidity.
Increasing Acidity
Between the mid-nineteenth century and World War II, the Industrial Rev-
olution led to a tremendous increase in coal burning and metal ore process-
ing in both Europe and North America. The combustion of coal, which
contains an average of 1.5 percent sulfur by weight, and the smelting of
metal sulfides released opaque plumes of smoke and sulfur dioxide from
short chimneys into the atmosphere.
Copper, nickel, and zinc smelters fumigated nearby landscapes with
sulfur dioxide and heavy metals. One of the world’s largest nickel smelt-
ers, located in Sudbury, Ontario, Canada, began operation in 1890 and by
1960 was pouring 2.6 million tons of sulfur dioxide per year into the atmo-
sphere. By 1970 the environmental damage extended to 72,000 hectares of
injured vegetation, lakes, and soils surrounding the site; within this area
17,000 hectares were barren. The land was devastated not only by acid de-
1
Acid deposition
Activists blame coal-burning

power plants and factory
emissions for acid rain problems.
After being emitted by large,
stationary sources, especially
those that have very high
smokestacks, pollutants can travel
thousands of kilometers in the
atmosphere. Those that are
transformed into sulfuric and
nitric acid aerosols are
incorporated into precipitation,
which eventually makes contact
with the earth’s surface.
(PhotoDisc)
position but also by the accumulation of toxic metals in the soil, the clear-
cutting of forested areas for fuel, and soil erosion caused by wind, water,
and frost heave.
In urban areas, high concentrations of sulfur corroded metal and accel-
erated the erosion of stone structures. During the winter, the added emis-
sions from home heating and stagnant weather conditions caused severe
air pollution episodes characterized by sulfuric acid fogs and thick, black
soot. In 1952 a four-day air pollution episode in London, England, killed an
estimated four thousand people.
After World War II, large coal-burning utilities in Western Europe and
the United States built their plants with particulate control devices and tall
stacks (higher than 100 meters) to improve the local air quality. Huge in-
dustrial facilities throughout Eastern Europe and the Soviet Union oper-
ated without air pollution controls for most of the twentieth century. The
tall stacks increased the dispersion and transport of air pollutants from
tens to hundreds of kilometers. Worldwide emissions of sulfur dioxide in-
creased; in the United States emissions climbed from 18 million tons in
1940 to a peak of 28 million tons in 1970. Acid deposition evolved into an
interstate and even an international problem.
2
Acid deposition
In major cities, exhaust from automobiles combined with power plant

and industrial emissions to create a choking, acrid smog of ozone, and ni-
tric and organic acids formed by photochemical processes. The rapid dete-
rioration of air quality in cities, with the attendant health and environmen-
tal consequences, spurred the passage of the U.S. Clean Air Act (CAA) of
1963, which was amended and expanded in 1970, 1977, and 1990. Each
amendment to the CAA brought new requirements for air pollution con-
trols.
Effects on Aquatic Ecosystems

The nature and extent of the environmental impact of acid deposition are
in dispute. Landscapes or surface waters impoverished by limestone or
acid-buffering soils are more sensitive to acid deposition. Regions that are
both sensitive and exposed to acid deposition include the eastern United
States, southeastern Canada, southern Sweden and Norway, central and
Eastern Europe, the United Kingdom, southeastern China, and the north-
ern tip of South America. Scientists hypothesize that within these regions
acid rain disrupts aquatic ecosystems and contributes to forest decline.
In southern Norway, for example, fish have been virtually extinct since
the late 1970’s in four-fifths of the lakes and streams in an area of 2 mil-
Acid Precipitation pH Scale

Natural
background
Seawater Pure water precipitation Citric juices
8 7 6 5 4 3 2 1 pH
Alkaline Acid
Most surface fresh waters
Increasing risk
to organisms
Acid precipitation,
eastern U.S., Scandinavia
Acid precipitation,
western U.S.
Acidified lakes and streams,
northeastern U.S., Scandinavia
Source: Adapted from John Harte, “Acid Rain,” in The Energy-Environment Connection,
edited by Jack M. Hollander, 1992.
Note: The acid precipitation pH ranges given correspond to volume-weighted annual
averages of weekly samples.
3
Acid deposition
lion hectares. Records and long-term monitoring showed that the decline
of fish populations began in the early twentieth century with dramatic
losses in the 1950’s. A strong correlation has been found between fish ex-
tinction and lake acidity. Researchers have also found that the diversity of
not only fish but also phytoplankton, zooplankton, invertebrates, and am-
phibian species diminishes by more than 50 percent as lake water pH
drops from 6.0 to 5.0. Below pH 5.6, aluminum released from lake sedi-
ments or leached from the surrounding soils interferes with gas and ion ex-
change in fish gills and can be toxic to aquatic life. Below pH 4.0, no fish
survive.
In the United States, 11 percent of the lakes in the Adirondack Moun-
tains in New York are too acidic to sustain fish life. Much controversy sur-
rounds the claim that these lakes were acidified by plumes of air pollutants
carried by prevailing winds from the Ohio River Valley. Like the lakes in
Norway, the Adirondack lakes have low acid-neutralizing capacity. Fish
declines that began in the early twentieth century and continued through
the 1980’s corresponded to reductions in pH. Fish kills often followed
spring snowmelt, which filled the waterways with acid accumulated in
winter precipitation. Historical records, field observations, and laboratory
experiments contradict arguments that overfishing, disease, or water pol-
lution killed the fish.
Effects on Forests and Cities

In areas exposed to acid rain, dead and dying trees stand as symbols of en-
vironmental change. In Germany the term Waldsterben, or forest death, is
used to describe the rapid declines of Norway spruce, Scotch pine, and sil-
ver fir trees in the early 1980’s, followed by beech and oak trees in the late
1980’s, especially at high elevations in the Black and Bavarian Forests. At
higher altitudes, clouds frequently shroud mountain peaks, bathing the
forest canopy in a mist of heavy metals, and sulfuric and nitric acids. Un-
der drought conditions, invisible plumes of ozone from sources hundreds
of kilometers distant intercept the mountain slopes. Several forests within
the United States are likewise affected, including the forests of ponderosa
pine in the San Bernardino Mountains of California, balsam fir in the
Smokey Mountains of North Carolina and Tennessee, and red spruce in the
Green and White Mountains of New England.
After more than one decade of intensive field and laboratory investiga-
tions of forest decline in North America and Europe, the link between dead
trees and acid deposition remained little more than circumstantial. Labora-
tory experiments often showed that acid rain had no effect, or even a fertil-
izing effect, on trees. Changes in foliage color, size, and shape; destruction
4
Acid deposition
of fine roots and associated fungi; and stunted growth are symptoms of
tree stress. Many researchers attribute these symptoms and forest decline
to the interactions of acid precipitation, ozone, excessive nitrogen deposi-
tion, land management practices, climate change, drought, and pestilence.
Ambient air concentrations of sulfur dioxide and nitrogen oxides are
typically higher in major cities, a result of the high density of emission
sources. The acids they form accelerate the weathering of exposed stone,
brick, concrete, glass, metal, and paint. For example, the calcite in lime-
stone and marble reacts with water and sulfuric acid to form gypsum (cal-
cium sulfate). The gypsum washes off stone with rain or, if eaves protect
the stone, accumulates as a soot-darkened crust. The acid-induced weath-
ering obscures the details of elaborate carvings on medieval cathedrals, an-
cient Greek columns, and Mayan ruins at alarming rates. Graffiti, pigeon
excrement, and the growth of bacteria and fungi on rock surfaces may
compound the damage.
Prevention Efforts
In the United States, the Acidic Deposition Control Program, Title IV of the
Clean Air Act amendments of 1990, directed the Environmental Protection
Agency (EPA) to reduce the adverse effects of acid rain. Public law man-
dated that the United States achieve 40 percent and 10 percent annual re-
ductions in sulfur dioxide and nitrogen dioxide emissions, respectively, by
the year 2000 from a 1980 base. The National Acid Precipitation Assess-
ment Program coordinates interagency acid deposition monitoring and re-
search, and assesses the cost, benefits, and effectiveness of acid deposition
control strategies. This echoes the 1985 30 Percent Protocol of the Conven-
tion on Long-Range Transboundary Air Pollution. Twenty-one nations
signed the protocol, thereby agreeing to reduce sulfur dioxide emissions 30
percent from 1980 levels by 1993.
Strategies to reduce acid deposition in the United States target large
electric utilities responsible for 70 percent of the sulfur dioxide and 30 per-
cent of the nitrogen oxide emissions. Utilities participate in a novel market-
based emission allowance trading and banking system that permits great
flexibility in controlling sulfur dioxide emissions. For example, a utility
may choose to remove sulfur from coal by cleaning it, burn a cleaner fuel
such as natural gas, or install a gas desulfurization system to reduce emis-
sions. The $6 billion international Clean Coal Technology Demonstration
Program, funded by governments and private industries, continues to de-
velop technologies—such as catalytic conversion of nitrogen oxides to in-
ert nitrogen—to radically decrease emissions of acid gases from coal-fired
power plants.
5
Acid deposition
Computer models of acid deposition in the northeastern United States

predicted that a 50 percent reduction in sulfur dioxide emissions would
decrease sulfur deposition by 44 to 48 percent. Between 1980 and 1996, U.S.
electric utilities lowered annual emissions of sulfur dioxide by 30 percent
from 17.5 million tons and nitrogen oxides by 14 percent from 7 million
tons. Average ambient air concentrations of sulfur dioxide decreased 37
percent and nitrogen oxide concentrations 10 percent between 1987 and
1996. However, a similar trend in sulfate and nitrate deposition has not
been observed.
Noreen D. Poor
See also: Lakes and limnology; Marine biomes; Ocean pollution and oil
spills; Reefs.
Sources for Further Study

Adriano, D. C., and A. H. Johnson, eds. Biological and Ecological Effects. Vol.
2 in Acidic Precipitation. New York: Springer-Verlag, 1989.
Ahrens, C. Donald. Meteorology Today. 6th ed. Pacific Grove, Calif.: Brooks/
Cole, 2000.
Canter, Larry W. Acid Rain and Dry Deposition. Chelsea, Mich.: Lewis, 1989.
Ellerman, A. Denny, et al. Markets for Clean Air: The U.S. Acid Rain Program.
New York, N.Y.: Cambridge University Press, 2000. Illustrated.
Gunn, John M., ed. Restoration and Recovery of an Industrial Region: Progress
in Restoring the Smelter-Damaged Landscape near Sudbury, Canada. New
York : Springer-Verlag, 1995.
Lutgens, Frederick K., and Edward J. Tarbuck. The Atmosphere. 8th ed. Up-
per Saddle River, N.J.: Prentice Hall, 2001.
U.S. Geological Survey. Acid Rain and Our Nation’s Capital. Author, 1997.
6
ADAPTATIONS AND THEIR
MECHANISMS
Types of ecology: Evolutionary ecology; Physiological ecology
Adaptations are structures, physiological mechanisms, or behaviors that are shaped

by the environment and enable organisms to cope with specific environmental con-
ditions. Studying adaptations helps scientists understand how organisms live
with environmental constraints and allows them to examine the mechanisms of
evolution.
W hy so many species exist is one of the most intriguing questions of

ecology. The study of adaptations offers an explanation. Because
there are many ways to cope with the environment, and because natural
selection has guided the course of evolutionary change for billions of
years, the vast variety of species existing on the earth today is simply an ex-
tremely complicated variation on the theme of survival. Many of the fea-
tures that are most interesting and beautiful in biology are adaptations.
Adaptations are the result of long evolutionary processes in which suc-
ceeding generations of organisms become better able to live in their envi-
ronments. Specialized structures, physiological processes, and behaviors
are all adaptations when they allow organisms to cope successfully with
the special features of their environments. Adaptations ensure that indi-
viduals in populations will reproduce and leave well-adapted offspring,
thus ensuring the survival of the species.
Mutation and Natural Selection

Adaptations arise through mutations—heritable changes in an organism’s
genetic material. These rare events are usually harmful, but occasionally
they give specific survival advantages to the mutated organism and its off-
spring. When certain individuals in a population possess advantageous
mutations, they are better able to cope with their specific environmental
conditions and, as a result, will contribute more offspring to future genera-
tions compared with those individuals in the population that lack the mu-
tation. Over time, the number of individuals that have the advantageous
mutation will increase in the population at the expense of those that do not
have it. Individuals with an advantageous mutation are said to have a
higher “fitness” than those without it, because they tend to have compara-
tively higher survival and reproductive rates. This is natural selection.
7
Adaptations and their mechanisms
Over very long periods of time, evolution by natural selection results in

increasingly better adaptations to environmental circumstances. Natural
selection is the primary mechanism of evolutionary change, and it is the
force that either favors or selects against mutations. Although natural se-
lection acts on individuals, a population gradually changes as those with
adaptations become better represented in the total population. Predaceous
fish, for example, which rely on speed to pursue and overtake prey, would
benefit from specific adaptations that would increase their swimming
speed. Therefore, mutations causing a sleeker and more hydrodynamically
efficient form would be beneficial to the fish predator. Such changes would
be adaptations if they resulted in improved predation success, diet, and re-
productive success, compared with slower members of the population.
Natural selection would favor the mutations because they confer specific
survival advantages to those that carry the mutations and impose limita-
tions on those lacking these advantages. Thus, those individuals with spe-
cial adaptations for speed would have a competitive advantage over
slower-swimming individuals. These attributes would be passed to their
more numerous offspring and, in evolutionary time, speed and hydrody-
namic efficiency would increase in the population.
General vs. Specific Adapations

Adaptations can be general or highly specific. General adaptations define
broad groups of organisms whose general lifestyle is similar. For example,
mammals are homeothermic, provide care for their young, and have many
other adaptations in common. At the species level, however, adaptations
are more specific and give narrow definition to those organisms that are
more closely related to one another. Slight variations in a single character-
istic, such as bill size in the seed-eating Galápagos finches, are adaptive in
that they enhance the survival of several closely related species. An under-
standing of how adaptations function to make species distinct also fur-
thers the knowledge of how species are related to one another.
Although natural selection serves as the instrument of change in shap-
ing organisms to very specific environmental features, highly specific ad-
aptations may ultimately be a disadvantage. Adaptations that are special-
ized may not allow sufficient flexibility (generalization) for survival in
changing environmental conditions. The degree of adaptative specializa-
tion is ultimately controlled by the nature of the environment. Environ-
ments, such as the tropics, that have predictable, uniform climates and
have had long, uninterrupted periods of climatic stability are biologically
complex and have high species diversity. Scientists generally believe that
this diversity results, in part, from complex competition for resources and
8
from intense predator-prey interactions. Because of these factors, many

narrowly specialized adaptations have evolved when environmental sta-
bility and predictability prevail. By contrast, harsh physical environments
with unpredictable or erratic climates seem to favor organisms with gen-
eral adaptations, or adaptations that allow flexibility. Regardless of the en-
vironment type, organisms with both general and specific adaptations ex-
ist because both types of adaptation enhance survivorship under different
environmental circumstances.
Structural Adaptations
Structural adaptations are parts of organisms that enhance their survival
ability. Camouflage, enabling organisms to hide from predators or their
prey; specialized mouth parts that allow organisms to feed on specific food
sources; forms of appendages, such as legs, fins, or webbed toes, that allow
efficient movement; protective spines that make it difficult for the organ-
ism to be eaten—these are all structural adaptations. These adaptations en-
hance survival because they assist individuals in dealing with the rigors of
the physical environment, obtaining nourishment, competing with others,
or hiding from or confusing predators.
Metabolic and Physiological Adaptations

Metabolism is the sum of all chemical reactions taking place in an organ-
ism, whereas physiology consists of the processes involved in an organism
carrying out its function. Physiological adaptations are changes in the me-
tabolism or physiology of organisms, giving them specific advantages for
a given set of environmental circumstances. Because organisms must cope
with the rigors of their physical environments, physiological adaptations
for temperature regulation, water conservation, varying metabolic rate,
and dormancy or hibernation allow organisms to adjust to the physical en-
vironment or respond to changing environmental conditions.
Desert environments, for example, pose a special set of problems for or-
ganisms. Hot, dry environments require physiological mechanisms that
enable organisms to conserve water and resist prolonged periods of high
temperature. Highly efficient kidneys and other excretory organs that as-
sist organisms in retaining water are physiological adaptations related to
the metabolisms of desert organisms. The kangaroo rat is a desert rodent
extremely well adapted to its habitat. Kangaroo rats do not drink, but
rather can obtain all of their water from the seeds they eat. They produce
highly concentrated urine and feces with very low water content.
Adaptation to a specific temperature range is also an important physio-
logical adaptation. Organisms cannot live in environments with tempera-
9
tures beyond their range of thermal tolerance, but some organisms are
adapted to warmer and others to colder environments. Metabolic response
to temperature is quite variable among animals, but most animals are
either homeothermic (warm-blooded) or poikilothermic (cold-blooded).
Homeotherms maintain constant body temperatures at specific tempera-
ture ranges. Although a homeotherm’s metabolic heat production is con-
stant when the organism is at rest and when environmental temperature is
constant, strenuous exercise produces excess heat that must be dissipated
into the environment, or overheating and death will result. Physiological
adaptations that enable homeotherms to rid their bodies of heat are the
ability to increase blood flow to the skin’s surface, sweating, and panting,
all of which promote heat loss to the atmosphere.
Behavioral Adaptations
Behavioral adaptations allow organisms to respond appropriately to vari-
ous environmental stimuli. Actions taken in response to various stimuli
are adaptive if they enhance survivorship. Migrations are behavioral ad-
aptations because they ensure adequate food supplies or the avoidance of
adverse environmental conditions. Courtship rituals that help in species
recognition prior to mating, reflex and startle reactions allowing for quick
retreats from danger, and social behavior that fosters specialization and co-
operation for group survival are behavioral adaptations.
Coevolution
Because organisms must also respond and adapt to an environment filled
with other organisms—including potential predators and competitors—
adaptations that minimize the negative effects of biological interactions
are favored by natural selection. Many times the interaction between spe-
cies is so close that each species strongly influences the others in the inter-
action and serves as the selective force causing change. Under these cir-
cumstances, species evolve together in a process called coevolution. The
adaptations resulting from coevolution have a common survival value to
all the species involved in the interaction. The coevolution of flowers and
their pollinators is a classic example of these tight associations and their re-
sulting adaptations.
The Peppered Moth

A classic example of recent evolutionary change and adaptation comes
from England. The peppered moth with a mottled gray color, is well
adapted to resting quietly on pale tree bark, with which it blends nicely.
This adaptive coloration (camouflage) enhanced the moth’s survival be-
10
cause the moths could remain largely undetected by predators during day-
light hours. Between 1850 and 1950, however, industrialization near urban
centers blackened tree trunks with soot, making the gray form disad-
vantageous, as it stood out on the contrasting background. During this
period, the gray moths began to disappear from industrial areas, but a
black-colored variant, previously rare, became increasingly common in the
population. These circumstances made it possible for scientists to test
whether the peppered moth’s camouflage was adaptive.
In a simple experiment, moths were raised in the laboratory, and equal
numbers of gray and black moths were released in both industrial and un-
polluted rural areas. Sometime later, only half of the gray-colored moths
could be recovered from the industrial sites, while only half of the black
forms could be recovered from the rural sites, compared with the total
number released. These results enabled the scientists to conclude that in-
creased predation on the gray moths in industrial areas led to a greater
fitness of the black moths, so the frequency of black moths increased in
the population. The reverse was true at the rural sites. This is the first well-
documented case of natural selection causing evolutionary change, and it
illustrates the adaptive significance of camouflage.
Robert W. Paul
See also: Adaptive radiation; Biodiversity; Biogeography; Camouflage;

Clines, hybrid zones, and introgression; Coevolution; Defense mecha-
nisms; Evolution: history; Isolating mechanisms; Natural selection; Non-
random mating, genetic drift, and mutation; Population genetics; Punctu-
ated equilibrium vs. gradualism; Speciation; Species loss; Trophic levels
and ecological niches.

Birkhead, Mike, and Tim Birkhead. The Survival Factor. New York: Facts on
File, 1990.
Brandon, Robert N. Adaptation and Environment. Princeton, N.J.: Princeton
University Press, 1990.
Gould, Stephen J. Ever Since Darwin. New York: W. W. Norton, 1977.
Ricklefs, Robert E. Ecology. 4th ed. New York: W. H. Freeman, 1999.
Rose, Michael R., and George V. Lauder, eds. Adaptation. San Diego, Calif.:
Academic Press, 1996.
Weibel, Ewald R. Symmorphosis: On Form and Function in Shaping Life. Cam-
bridge, Mass.: Harvard University Press, 2000.
Whitfield, Philip. From So Simple a Beginning: The Book of Evolution. New
York: Macmillan, 1995.
11
ADAPTIVE RADIATION
Types of ecology: Evolutionary ecology; Population ecology; Speciation
In adaptive radiation, numerous species evolve from a common ancestor intro-

duced into an environment with diverse ecological niches. The progeny evolve ge-
netically into customized variations of themselves, each adapting to survive in a
particular niche.
I n 1898 Henry F. Osborn identified and developed the concept of adap-

tive radiation, whereby different forms of a species evolve, quickly in
evolutionary terms, from a common ancestor. According to the principles
of natural selection, organisms that are the best adapted (most fit) to com-
pete will live to reproduce and pass their successful traits on to their off-
spring. The process of adaptive radiation illustrates one way in which nat-
ural selection can operate when members of one population of a species
are cut off from another or migrate to a different environment that is iso-
lated from the first. Such isolation can occur from one patch of plantings to
another, from one mountaintop or hillside to another, from pond to pond,
or from island to island. Faced with different environments, the group will
diverge from the original population and in time become different enough
to form a new species.
Divergent Populations and Speciation

In a divergent population, the relative numbers of one form of allele (char-
acteristic) decrease, while the relative numbers of a different allele in-
crease. New environmental pressures will select for favorable alleles that
may not have been favored in the old environment. Over successive gener-
ations, therefore, a new gene created by random mutation may replace the
original form of the gene if, for example, the trait encoded by that gene al-
lows the divergent group to cope better with environmental factors, such
as food sources, predators, or temperature. The result in the long term is
that molecular material that forms genes, deoxyribonucleic acid (DNA),
changes sufficiently through the growth of divergent populations to allow
new generations to become significantly different from the original popu-
lation. In time, the new population is unable to reproduce with members of
the original population and becomes a new species.
Adaptive Radiation of Animals

Adaptive radiation occurs dramatically when a species migrates from one
12
Adaptive radiation
landmass to another. This may occur between islands or between conti-

nents and islands. A classic example of adaptive radiation is the evolution
of finches noted by Charles Darwin during his trips to the Galápagos Is-
lands off the west coast of South America. Several species of plants and an-
imals had migrated to these islands from the South American mainland by
means of flight, wind, ocean debris, or other means of transport. Finches
from the mainland—perhaps aided by winds—settled on fifteen of the is-
lands in the Galápagos group and began to adapt to the various unoccu-
pied ecological niches on those islands, which differed. Over several gen-
erations, natural selection favored a variety of finch species with beaks
adapted for the different types of foods available on the different islands.
As a result, several species of different finches evolved, roughly simulta-
neously, on these islands.
A more recent example of adaptive radiation in its early stages has
taken place in an original population of brown bears. The brown bear can
be found throughout the Northern Hemisphere, ranging from the decidu-
ous forests up into the tundra. During one of the glacier periods, a small
population of the brown bear was separated from the main group; accord-
ing to fossil evidence, this small population, under selection pressure from
the Arctic environment, evolved into the polar bear. Although brown bears
are classified as carnivores, their diets are mostly vegetarian, with occa-
sional fish and small animals as supplements. On the other hand, the polar
bear is mostly carnivorous. Besides its white coat, the polar bear is different
from the brown bear in many ways, including its streamlined head and
shoulders and the stiff bristles that cover the soles of its feet, which provide
traction and insulation, enabling it to walk on ice.
Adaptive Radiation of Plants

Although plants seem unable to “migrate” as birds and other animals do,
adaptive radiation occurs in the plant world as well. In the Hawaiian Is-
lands, for example, twenty-eight species of the Asteraceae family are known
together as the Hawaiian silversword alliance. The entire group appears to
be traceable to one ancestor, thought to have arrived on the island of Kauai
from western North America. The silverswords—which compose three
genera, Argyroxiphium, Dubautia, and Wilkesia—have since evolved into
twenty-eight species, and this speciation came about due to major ecologi-
cal shifts. These plants are therefore prime examples of adaptive radiation.
Within the silversword alliance, different species have adapted to
widely varying ecosystems found throughout the islands. Argyroxiphium
sandwicense, for example, is endemic to the island of Maui and grows at
high elevations from 6,890 to 9,843 feet (2,100-3,000 meters) on the dry, al-
13
Adaptive radiation
pine slopes of the volcano Haleakala. This species has succulent leaves
covered with silver hairs. It is thought that the hairs lessen the pace of
evaporative moisture loss and protect the leaves from the sun. In contrast,
species of the genus Dubautia that grow in wet, shady forests have large
leaves that lack hairs.
Despite their “customized” physiologies, the silverswords that have
evolved in Hawaii are all closely related to one another, so much so that
any two can hybridize. Studies of the silverswords have provided what ge-
neticist Michael Purugganan called a “genetic snapshot of plant evolu-
tion.”
Jon P. Shoemaker, updated by Bryan Ness
See also: Adaptations and their mechanisms; Biodiversity; Biogeography;

Clines, hybrid zones, and introgression; Coevolution; Competition; Con-
vergence and divergence; Evolution: definition and theories; Evolution:
history; Evolution of plants and climates; Extinctions and evolutionary ex-
plosions; Gene flow; Genetic diversity; Genetic drift; Isolating mecha-
nisms; Natural selection; Nonrandom mating, genetic drift, and mutation;
Population genetics; Punctuated equilibrium vs. gradualism; Speciation;
Trophic levels and ecological niches.

Givnish, Thomas J., and Kenneth J. Sytsma, eds. Molecular Evolution and
Adaptive Radiation. New York: Cambridge University Press, 2000.
Robichaux, Robert, et al. “’Radiating’ Plants.” Endangered Species Bulletin
Update, March/April, 1999, S4-S5.
Schluter, Dolph. The Ecology of Adaptive Radiation. Oxford, England: Oxford
14
ALLELOPATHY
Types of ecology: Chemical ecology; Community ecology
Allelopathy refers to all the biochemical interactions between species, including

microorganisms.
F or an allelopathic interaction to occur, chemicals must be released into

the environment by one organism that will affect the growth of an-
other. In this way allelopathy differs from competition, which involves re-
moval of some factor from the environment that is shared with other or-
ganisms.
Allelopathy was recognized as early as Theophrastus (300 b.c.e.), who
pointed out that chick pea plants destroy weeds growing around them.
Methods of Action
A variety of different allelochemicals are produced by plants, usually as
secondary metabolites that do not have a specific function in the growth
and development of the host plant but that do affect the growth of other
plants. Originally plant physiologists thought these secondary products
were simply metabolic wastes that plants had to store because they do not
have an excretory system as animals do. Their various functions are now
beginning to be understood.
One class of allelochemicals, coumarins, block or slow cell division in
the affected plant, particularly in root cells. In this way growth of compet-
ing plants is inhibited, and seed germination can be prevented. Several
kinds of allelochemicals, including flavonoids, phenolics, and tannins,
suppress or alter hormone production or activity in competing plants.
Other chemicals, including terpenes and certain antibiotics, alter mem-
brane permeability of host cells, making them either leaky or imperme-
able. In some cases, membrane uptake can be enhanced, particularly for
micronutrients in low concentration in the soil. Finally, a variety of
allelochemicals have both positive and negative effects on metabolic activ-
ity of the affected plant.
Allelopathy in Agriculture
Most of the negative effects of weeds on crop plants have been attributed to
competition; however, experiments using weed extracts have demon-
strated that many weeds produce allelochemicals. Similarly, some crop
plants are allelopathic to others and themselves, including wheat, corn,
15
Allelopathy
Some plant species, including peach trees, release chemicals into the soil that inhibit
the growth of other plants that might otherwise compete with them. (PhotoDisc)
and rice. In these cases the residues of one year’s crop can interfere with
crop growth in subsequent years. This is increasingly important for farm-
ers to consider who are incorporating low-tillage methods to reduce soil
erosion. To minimize these effects, some of the traditional techniques of
cover cropping, companion cropping, and crop rotation must be em-
ployed. Known allelopaths are also beginning to be used as biological con-
trol agents to manage invasive and weedy plant species.
Allelopathy in Nature
Several tree species, including black walnut, black locust, and various
pines, are known to produce allelochemicals that inhibit the growth of
understory species. In some cases this is a result of drip from the foliage or
leachate from fallen leaves and fruit. In other cases, roots secrete
allelochemicals that kill seedlings of other plants. Bracken fern (Pteridium
aquilinum) is known to affect the growth of many other plants.
Marshall D. Sundberg
See also: Animal-plant interactions; Biological invasions; Coevolution;

Communities: ecosystem interactions; Competition; Defense mechanisms;
16
Allelopathy
Invasive plants; Metabolites; Poisonous plants; Trophic levels and ecologi-

cal niches.

Moore, Randy, W. Dennis Clark, and Darrell S. Vodopich. Botany. 2d ed.
New York: McGraw-Hill, 1998.
Rice, Elroy L. Allelopathy. 2d ed. Orlando, Fla.: Academic Press, 1984.
17
ALTRUISM
Type of ecology: Behavioral ecology
Altruistic behavior involves an individual’s sacrifice of self in order to help others.

In some animals, altruism appears to be genetically determined.
T hose who study animal behavior (ethology) have observed that on oc-
casion individuals act altruistically. In other words, they appear vol-
untarily to put the needs of their group or of another individual ahead of
their own needs. According to some scientists, there are examples in nature
where a particular species might not have survived had there not been sac-
rifice by some on behalf of the many. One important question is whether
this so-called altruism has been a matter of voluntary choice or whether it
has occurred as a part of the selection process, making it, therefore, an in-
voluntary response.
Group Formation
Of interest to a wide group, including psychologists, sociologists, philoso-
phers, and political scientists, are the questions of whether altruism is de-
sirable behavior—perhaps even to the exclusion of egoism—and whether
altruism may be necessary for human survival. Some wonder whether
such behavior is necessary, whether it can be learned, and whether hu-
mans will voluntarily choose to learn it. Biologists and geneticists have
been left the problem of determining, if possible, whether the tendency for
altruism is inherited or learned behavior. Unfortunately for scientists, the
study of human beings in social groups in the wild is virtually impossible.
However, the study of animal behavior, primarily in native habitats, has
provided some insight, although it must be recognized that different spe-
cies have solved problems of survival in different ways.
Animals of the same species are bound to consort, if only for mating
purposes. Most species are, in fact, found to live in groups, not only for
purposes of reproduction but also because sources of food attract individ-
uals to the same places and because congregation provides better protec-
tion from predators. It is common in nature for groups to form because
their individual members have the same physical needs, and such groups
may stay together as long as the needs of those individuals can be met. This
does not necessarily mean that there exists in the group any loyalty or even
any recognition of individuals as members of the group. In more highly
developed societies, however, groups such as families or tribes develop.
18
Altruism
Offspring and Reproduction

In animal life, two or more adults and their offspring often form close
bonds and tend to exclude those who are not related. Each recognizes the
others as being members, and membership is restricted to those who are
among the founders or who are born into the smaller group and who con-
form in recognizable ways to the norms of the group. Hierarchy or rank is
recognized, and often there is a division of labor within the group.
It has been demonstrated that species that spend a large amount of time
providing for their young tend to have developed higher social orders.
Humans, for example, must care for their young much longer, before they
are able to become independent, than must many of the lower forms of ani-
mal life. Humans are aware of a bond that almost always exists between
parent and child and of the spirit of mutual support and a cooperation that
may exist even in the extended family. Cooperative behavior within such a
familial group may be considered to benefit all members. Because such be-
havior is not consistent, however—there are times when such bonds do not
exist and when families are not cooperative—such behavior cannot neces-
sarily be attributed to predisposition. Some have argued that in primitive
animal societies, so-called altruism may have evolved of necessity in order
to achieve reproductive success, but that in human society there may be no
evolutionary explanation for the phenomenon. Indeed, it could be argued
that pure altruism, for humans, might be self-defeating and therefore un-
likely to have developed as an inherited trait.
Evidence has been gathered in the study of some Hymenoptera (the or-
der of insects that includes bees, wasps, ants, sawflies, and other colony-
forming insects) that certain members of the population forage for the
group while others lay eggs and remain at the nest to guard them. Where
such behavior has evolved, through the necessity of feeding and protect-
ing those that will propagate their kind, the foragers may be labeled altru-
istic: They have sacrificed their own reproductive possibilities for survival
of the group. Some have questioned whether this phenomenon can truly
be labeled altruism, however, because the donor appears to be “pro-
grammed” to perform such behaviors rather than having a choice not to
perform the behaviors (conscious purpose is very difficult to assess in ani-
mals). Moreover, some researchers wonder how the traits that favor altru-
istic behavior can survive and become dominant in a group if those having
the traits deemed desirable are not allowed to reproduce. With the use of
mathematical models, it has been demonstrated that such traits can be pre-
served only within the family unit.
Among close relatives, the traits appear with enough strength that they
will be reproduced in a greater concentration, thereby compensating for
19
Altruism
the loss suffered by the sacrifice of the donors. This phenomenon has been
referred to as kin selection, because it occurs in groups that have strong rec-
ognition of membership—to the extent that there exists aggressive defense
against intruders, even of the same species. Discrimination against outsid-
ers is an important facet of altruism of this type. The willingness of an indi-
vidual to provide for others at the expense of its own interests diminishes
as the degree of relatedness decreases.
Reciprocal Sacrifice
Most parental behavior would not be labeled altruistic, since it is in the in-
terest of the parent to care for the offspring in order to ensure the survival
of the parent’s genes. Of perhaps more interest than what happens among
closely related members of a group and even between parent and offspring
is the question of what motivates sacrifice on the part of an individual
when no close relationship with the recipient exists—for example, a male
animal coming to the rescue of an unrelated male animal who is being at-
tacked by a third male of the same species.
One theory maintains that these acts of personal sacrifice are performed
on the chance that reciprocal sacrifice may occur at some future time.
Whether this type of altruism can occur through natural selection, which
acts through individuals, is an interesting question. Models have shown
that in a population where individuals are likely to encounter and recog-
nize one another on a frequent basis, it is possible that reciprocal exchanges
can take place. Individual A might be the donor on the first encounter, in-
dividual B on the second. This theory requires that the two must have
a high probability of subsequent encounters and that the tendency for al-
truism must already have been established through kin selection. Be-
cause animals are usually suspicious of strangers on first encounter, it is
necessary to speculate that in its beginning, altruism was a selected-for
trait in very small groups where strangers were not only nonhostile but
also likely to be relatives and likely to be met again. This type of behavior,
in which individuals act in a manner not to their own advantage and not in
order that their own genes or the genes of relatives will survive, is per-
formed, in theory, with some expectation of imagined reciprocal gain.
How this type of behavior has come about, however, is a matter requiring
further study.
Cultural Influences
Another question concerns how much culture is an influence on the devel-
opment of a hereditary tendency toward altruism. Some have suggested
that after generations and generations of cultural emphasis on the need for
20
Altruism
altruism, it might come to have a genetic basis. There is little hard evidence
that this would occur. On the other hand, humans have had a very rapid
cultural evolution, and it possible that they may have had strong genetic
propensities for altruism which have been culturally overlaid. Some argue
that biology and culture evolve simultaneously—that the culture is formed
as a result of the imposition of genetic factors while, at the same time, ge-
netic traits are evolving in response to cultural change. In order to under-
stand the source of altruism in humankind, one must study such behavior
in the context of many factors in human development—biological as well
as social, cultural, economic, and ecologic.
Studying Altruistic Behavior

Those investigating the sources of altruism usually begin with a thorough
understanding of whatever organism is the subject of the study. When the
insect or animal cannot be studied in the wild, the ethologist tries to simu-
late the important features of the natural habitat in a captive environment,
at least in the beginning. Models are devised, based on observable data;
formulas are employed; and projections are made, which provide a basis
for speculative argument when absolutes cannot be assured.
By observing, it is possible to determine whether various evidences of
altruism exist within a population. Altruism may be manifested in as sim-
ple a way as the sharing of food when there is a scarcity. In some popula-
tions, one might observe a division of labor in which some forfeit their re-
productive possibilities in order to care for the offspring of others. This
phenomenon introduces the question of how altruism can survive in a
population in which the genetic traits favoring the behavior are most evi-
dent in the individuals that do not reproduce themselves. It has been
shown that the tendency for altruism can be perpetuated only within the
family unit where the same genetic tendency exists to some degree in
members that engage in reproductive activity; this can be demonstrated by
a mathematical formula.
Each individual bears the inheritance coefficient or relatedness coeffi-
cient r. Offspring share with each parent an average of half of the genetic
traits of each (r = 1 2). Offspring share with each grandparent one-fourth of
the genetic traits of each of the older generation (r = 1 4); the same coefficient
exists with cousins. Were the altruists not to reproduce, it would be re-
quired, in order for the trait to be passed on, that the reproductive chances
of their siblings more than double or that the reproductive chances of their
cousins more than quadruple. For the sacrifice to be of value, the genetic
relationship must be close, according to the demonstration. The case has
been made that in societies having evolved according to this principle,
21
Altruism
there is a diminishing willingness to put the interests of others ahead of

one’s own as the degree of kinship decreases.
In societies where males are produced from unfertilized eggs and fe-
males from fertilized ones, female offspring of a mated pair have a high re-
latedness coefficient (r = 3 4). The altruists among the female siblings will
benefit more, regarding their genetic potential, by caring for their sisters
than for their own offspring, and it can again be observed that sacrifice is
more likely to be made on behalf of the member that is more closely re-
lated.
Voluntary vs. Involuntary Altruism

If altruism exists in nature, and if it has come about through natural selec-
tion, then one can argue that it must be a behavior with value. When apply-
ing the human connotation to the term altruism, however, one must con-
sider the role of choice in the manifestation of the behavior. Humans claim to
admire acts of unselfishness that are seemingly done with no expectation of
reward. The admiration would diminish or become nonexistent, however, if
there were to be proof that the act was performed because of some primitive
biological predisposition rather than because of a decision on the part of the
donor. Therefore, it is necessary to make the distinction, when discussing
the importance of altruism, as to whether one is referring to the acts of hu-
man beings that are performed in the face of emergency or tragedy, where
a sacrifice is made as a matter of choice, or whether the intent is to consider
altruism as it occurs in other creatures and seems to be involuntary.
In the case of nonhuman forms, altruism as an act of voluntary sacrifice
is infrequent—if indeed it exists at all. Altruism, however, as an act which
is dictated by genetics, is observable, and it has been shown to have been
necessary for the survival of certain species. Where animal societies have
formed in which some members of the society have spent their lives caring
for the offspring of others or performing other sacrificial behavior which
benefits the group, there can be little doubt that such altruism has been dic-
tated by nature for its own unique purposes.
Moreover, the fact that voluntary self-sacrifice on the part of human be-
ings does exist does not automatically make it desirable human behavior
any more than aggressive behavior is automatically undesirable. The case
can be made that both types of behavior are important. Perhaps the larger
question is when and under what circumstances certain types of human
behavior should be acceptable or desirable for the individual and for the
group, and, even more important, who is qualified to decide what type of
behavior is appropriate.
P. R. Lannert
22
Altruism
See also: Communication; Defense mechanisms; Displays; Ethology; Hier-

archies; Insect societies; Mammalian social systems; Mimicry; Pheromones;
Population genetics; Predation; Reproductive strategies; Territoriality and
aggression.

Boorman, Scott A., and Paul R. Levitt. The Genetics of Altruism. New York:
Academic Press, 1980.
Bradie, Michael. The Secret Chain: Evolution and Ethics. Albany: State Uni-
versity of New York Press, 1994.
Wright, Robert. Nonzero: The Logic of Human Destiny. New York: Pantheon
Books, 2000.
Zahn-Waxler, Carolyn, E. Mark Cummings, and Ronald Iannotti, eds. Al-
truism and Aggression: Biological and Social Origins. New York: Cam-
bridge University Press, 1986.
23
ANIMAL-PLANT INTERACTIONS
Types of ecology: Community ecology
The ways in which certain animals and plants interact have evolved in some cases
to make them interdependent for nutrition, respiration, reproduction, or other as-
pects of survival.
T he realm of ecology involves a systematic analysis of plant-animal in-

teractions through the considerations of nutrient flow in food chains
and food webs, exchange of such important gases as oxygen and carbon di-
oxide between plants and animals, and strategies of mutual survival be-
tween plant and animal species through the processes of pollination and
seed dispersal. Having the unique ability, by photosynthesis, to take car-
bon dioxide and incorporate it into organic molecules, green plants are
classified as ecological producers. Animals are classified as consumers,
taking the products of photosynthesis and chemically breaking them down
at the cellular level to produce energy for life activities. Carbon dioxide is a
waste product of this process. Given their respective status as producers
and consumers, plants and animals have over the ages formed many eco-
logical relationships.
Mutualism
Mutualism is an ecological interaction in which two different species of or-
ganisms beneficially reside together in close association, usually revolving
around nutritional needs. One such example is a small aquatic flatworm
that absorbs microscopic green algae into its tissues. The benefit to the ani-
mal is one of added food supply. The mutual adaptation is so complete that
the flatworm does not actively feed as an adult. The algae, in turn, receive
adequate supplies of nitrogen and carbon dioxide and are literally trans-
ported throughout tidal flats in marine habitats as the flatworm migrates,
thus exposing the algae to increased sunlight. This type of mutualism,
which verges on parasitism, is called symbiosis.
Coevolution
Coevolution is an evolutionary process wherein two organisms interact so
closely that thy evolve together in response to shared or antagonistic selec-
tion pressure. A classic example of coevolution involves the yucca plant
and a species of small, white moth (Tegitecula). The female moth collects
pollen grains from the stamen of one flower on the plant and transports
24
Animal-plant interactions
these pollen loads to the pistil of another flower, thereby ensuring cross-
pollination and fertilization. During this process, the moth will lay her
own fertilized eggs in the flowers’ undeveloped seed pods. The develop-
ing moth larvae have a secure residence for growth and a steady food sup-
ply. These larvae will rarely consume all the developing seeds; thus, both
species (plant and animal) benefit.
Although this example represents a mutually positive relationship be-
tween plants and animals, other interactions are more antagonistic. Preda-
tor-prey relationships between plants and animals are common. Insects
and larger herbivores consume large amounts of plant material. In re-
sponse to this selection pressure, many plants have evolved secondary me-
tabolites that make their tissues unpalatable, distasteful, or even poison-
ous. In response, herbivores have evolved ways to neutralize these plant
defenses.
Mimicry
In mimicry, an animal or plant has evolved structures or behavior patterns
that allow it to mimic either its surroundings or another organism as a de-
fensive or offensive strategy. Certain types of insects, such as the leaf-
hopper, walking stick, praying mantis, and katydid (a type of grasshopper),
often duplicate plant structures in environments ranging from tropical
rain forests to northern coniferous forests. Mimicry of their plant hosts af-
fords these insects protection from their own predators as well as camou-
flage that enables them to capture their own prey readily. Certain species of
ambush bugs and crab spiders have evolved coloration patterns that allow
them to hide within flower heads of such common plants as goldenrod, en-
abling them to ambush the insects that visit these flowers.
Nonsymbiotic Mutualism
In nonsymbiotic mutualism, plants and animals coevolve morphological
structures and behavior patterns by which they benefit each other but
without living physically together. This type of mutualism can be demon-
strated in the often unusual shapes, patterns, and colorations that more ad-
vanced flowering plants have developed to attract various insects, birds,
and mammals for pollination and seed dispersal purposes. Accessory
structures, called fruits, form around seeds and are usually tasty and
brightly marked to attract animals for seed dispersal. Although the fruits
themselves become biological bribes for animals to consume, often the
seeds within these fruits are not easily digested and thus pass through the
animals’ digestive tracts unharmed, sometimes great distances from the
parent plant. Some seeds must pass through the digestive plant of an ani-
25
mal to stimulate germination. Other types of seed dispersal mechanisms

involve the evolution of hooks, barbs, and sticky substances on seeds that
enable them to be easily transported by animal fur, feet, feathers, or beaks.
Such strategies of dispersal reduce competition between the parent plant
and its offspring.
Pollinators
Because structural specialization increases the possibility that a flower’s
pollen will be transferred to a plant of the same species, many plants have
evolved a vast array of scents, colors, and nutritional products to attract
pollinators. Not only does pollen include the plant’s sperm cells; it also rep-
resents a food reward. Another source of animal nutrition is a substance
called nectar, a sugar-rich fluid produced in specialized structures called
nectaries within the flower or on adjacent stems and leaves. Assorted
waxes and oils are also produced by plants to ensure plant-animal interac-
tions. As species of bees, flies, wasps, butterflies, and hawkmoths are at-
tracted to flower heads for these nutritional rewards, they unwittingly be-
come agents of pollination by transferring pollen from stamens to pistils.
Some flowers have evolved distinctive, unpleasant odors reminiscent
of rotting flesh or feces, thereby attracting carrion beetles and flesh flies in
search of places to reproduce and deposit their own fertilized eggs. As
these animals copulate, they often become agents of pollination for the
plant itself. Some tropical plants, such as orchids, even mimic a female bee,
wasp, or beetle, so that the insect’s male counterpart will attempt to mate
with them, thereby encouraging precise pollination.
Among birds, hummingbirds are the best examples of plant pollinators.
Various types of flowers with bright, red colors, tubular shapes, and
strong, sweet odors have evolved in tropical and temperate regions to take
advantage of hummingbirds’ long beaks and tongues as an aid to pollina-
tion. Because most mammals, such as small rodents and bats, do not detect
colors as well as bees and butterflies do, some flowers instead focus upon
the production of strong, fermenting, or fruitlike odors and abundant pol-
len rich in protein. In certain environments, bats and mice that are primar-
ily nocturnal have replaced day-flying insects and birds as pollinators.
Thomas C. Moon, updated by Bryan Ness
See also: Allelopathy; Coevolution; Communities: ecosystem interactions;

Communities: structure; Competition; Defense mechanisms; Food chains
and webs; Herbivores; Hierarchies; Lichens; Mycorrhizae; Omnivores;
Poisonous plants; Pollination; Predation; Symbiosis; Trophic levels and
ecological niches.
26

Abrahamson, Warren G., and Arthur E. Weis. Evolutionary Ecology Across
Three Trophic Levels: Goldenrods, Gallmakers, and Natural Enemies. Prince-
ton, N.J.: Princeton University Press, 1997.
Barth, Friedrich G. Insects and Flowers: The Biology of a Partnership. Prince-
Buchmann, Stephen L., and Gary Paul Nabhan. The Forgotten Pollinators.
Washington, D.C.: Island Press/Shearwater Books, 1997.
Dickerman, Carolyn. “Pollination: Strategies for Survival.” Ward’s Natural
Science Bulletin, Summer, 1986, 1-4.
Howe, Henry F., and Lynne C. Westley. Ecological Relationships of Plants and
Animals. New York: Oxford University Press, 1990.
John, D. M., S. J. Hawkins, and J. H. Price, eds. Plant-Animal Interactions in
the Marine Benthos. Oxford, England: Clarendon Press, 1992.
Lanner, Ronald M. Made for Each Other: A Symbiosis of Birds and Pines. New
York: Oxford University Press, 1996.
Meeuse, Bastian, and Sean Morris. The Sex Life of Flowers. New York: Facts
on File, 1984.
Price, Peter W., G. Wilson Fernandes, Thomas H. Lewinsohn, and Wood-
ruff W. Benson, eds. Plant-Animal Interactions: Evolutionary Ecology in
Tropical and Temperate Regions. New York: John Wiley & Sons, 1991.
Rudman, William B. “Solar-Powered Animals.” Natural History 96 (Octo-
ber, 1987): 50-53.
27
BALANCE OF NATURE
Types of ecology: Ecoenergetics; Theoretical ecology
The ecological concept of the balance of nature—a view that proposes that nature,
in its undisturbed state, is constant—has never been legitimized in science as ei-
ther a hypothesis or a theory. However, it laid the groundwork for the science of
ecology and persists as a designation for a healthy environment.
T he notion of the “balance of nature” has a deep history that dates back
to ancient times and has persisted into modern times. During the sci-
entific revolution in the seventeenth century, John Graunt, a merchant, an-
alyzed London’s baptismal and death records in 1662 and discovered
the balance in the sex ratio and the regularity of most causes of death (ex-
cluding epidemics). England’s chief justice, Sir Matthew Hale, was inter-
ested in Graunt’s discoveries, but he nevertheless decided that the human
population, in contrast to animal populations, must have steadily in-
creased throughout history. He surveyed the known causes of animal mor-
tality and in 1677 published the earliest explicit account of the balance of
nature.
English scientist Robert Hooke studied fossils and in 1665 concluded
that they represented the remains of plants and animals, some of which
were probably extinct. However, a clergyman-naturalist, John Ray, replied
that the extinction of species would contradict the wisdom of the ages, by
which he seems to have meant the balance of nature. Ray also studied the
hydrologic cycle, the geochemical cycle of water. Antoni van Leeuwen-
hoek, one of the first investigators to make biological studies with a micro-
scope, discovered that parasites are more prevalent than anyone had sus-
pected and that they are often detrimental or even fatal to their hosts.
Before that, it was commonly assumed that the relationship between host
and parasite was mutually beneficial.
Richard Bradley, a botanist and popularizer of natural history, pointed
out in 1718 that each species of plant has its own kind of insect and
that there are even different insects that eat the leaves and bark of a tree.
His book A Philosophical Account of the Works of Nature (1721) explored as-
pects of the balance of nature more thoroughly than had been done before.
Ray’s and Bradley’s books may have inspired the comment in Alexander
Pope’s Essay on Man (1733) that all species are so closely interdepen-
dent that the extinction of one would lead to the destruction of all living
nature.
28
Balance of nature
Toward a Science of Ecology

Swedish naturalist Carolus Linnaeus was an important protoecologist. His
essay Oeconomia Naturae (1749; The Economy of Nature, 1749) attempted to
organize the aspect of natural history dealing with the balance of nature,
but he realized that one must study not only ways that plants and animals
interact but also their habitats. He knew that while balance had to exist,
there occurred over time a succession of plants, beginning with a bare field
and ending with a forest. In Politia Naturae (1760; Governing Nature, 1760)
he discussed the checks on populations that prevent some species from be-
coming so numerous that they eliminate others. He noticed the competi-
tion among different species of plants in a meadow and concluded that
feeding insects kept them in check. French naturalist Comte de Buffon de-
veloped a dynamic perspective on the balance of nature from his studies
on rodents and their predators. Rodents can increase in numbers to plague
proportions, but then predators and climate reduce their numbers. Buffon
also suspected that humans had exterminated some large mammals, such
as mammoths and mastodons.
Later another Frenchman, Jean-Baptiste Lamarck, published his book
on evolution called Philosophie zoologique (1809; Zoological Philosophy, 1914),
which cast doubt on extinction by arguing that fossils only represent early
forms of living species: Mammoths and mastodons evolved into African
and Indian elephants. In developing this idea, he minimized the impor-
tance of competition in nature. An English opponent, the geologist Charles
Lyell, argued in 1833 that species do become extinct, primarily because of
competition among species. Charles Darwin was inspired by his own in-
vestigations during a long voyage around the world and by his reading of
the works of Linnaeus and Lyell. Darwin’s revolutionary book, On the Ori-
gin of Species by Means of Natural Selection (1859), argued an intermediate
position between Lamarck and Lyell: Species do evolve into different spe-
cies, but in the process, some species do indeed become extinct.
Darwin’s theory of evolution might have brought an end to the balance-
of-nature concept, but it did not. Instead, American zoologist Stephen A.
Forbes developed an evolutionary concept of the balance of nature in his
essay, “The Lake as a Microcosm” (1887). Although the reproductive rate
of aquatic species is enormous and the struggle for existence among them
is severe, “the little community secluded here is as prosperous as if its state
were one of profound and perpetual peace.” He emphasized the stabiliz-
ing effects of natural selection.
The Science of Ecology

The science of ecology became formally organized between the 1890’s and
29
Balance of nature
the 1910’s. One of its important organizing concepts was that of “biotic
communities.” An American plant ecologist, Frederic E. Clements, wrote a
large monograph titled Plant Succession (1916), in which he drew a mor-
phological and developmental analogy between organisms and plant com-
munities. Both the individual and the community have a life history dur-
ing which each changes its anatomy and physiology. This superorganismic
concept was an extreme version of the balance of nature that seemed plau-
sible as long as one believed that a biotic community was a real entity
rather than a convenient approximation of what one sees in a pond, a
meadow, or a forest. However, the studies of Henry A. Gleason in 1917 and
later indicated that plant species merely compete with one another in simi-
lar environments; he concluded that Clements’s superorganism was po-
etry, not science.
While the balance-of-nature concept was giving way to ecological hy-
potheses and theories, Rachel Carson decided that she could not argue her
case in Silent Spring (1962) without it. She admitted, “The balance of nature
is not a status quo; it is fluid, ever shifting, in a constant state of adjust-
ment.” Nevertheless, to her the concept represented a healthy environ-
ment, which humans could upset. Her usage of the phrase has persisted
within the environmental movement.
In 1972 English medical chemist James E. Lovelock developed a new
balance-of-nature idea, which he calls Gaia, named for a Greek earth god-
dess. His reasoning owed virtually nothing to previous balance-of-nature
notions that focused upon the interactions of plants and animals. His con-
cept emphasized the chemical cycles that flow from the earth to the waters,
atmosphere, and living organisms. He soon had the assistance of a zoolo-
gist named Lynn Margulis. Their studies convinced them that biogeo-
chemical cycles are not random, but exhibit homeostasis, just as some ani-
mals exhibit homeostasis in body heat and blood concentrations of various
substances. They believe that living beings, rather than inanimate forces,
mainly control the earth’s environment. In 1988 three scientific organiza-
tions sponsored a conference of 150 scientists from all over the world to
evaluate their ideas. Although science more or less understands how ho-
meostasis works when a brain within an animal controls it, no one has suc-
ceeded in satisfactorily explaining how homeostasis can work in a world
“system” that lacks a brain. The Gaia hypothesis is as untestable as were
earlier balance-of-nature concepts.
Frank N. Egerton
See also: Animal-plant interactions; Biomass related to energy; Biosphere

concept; Deep ecology; Ecology: definition; Ecosystems: definition and
30
Balance of nature
history; Evolution: history; Food chains and webs; Geochemical cycles;

Hydrologic cycle; Nutrient cycles; Trophic levels and ecological niches.

Arthur, Wallace. The Green Machine: Ecology and the Balance of Nature. Cam-
bridge, Mass.: Blackwell, 1990.
Egerton, Frank N. “Changing Concepts of the Balance of Nature.” Quar-
terly Review of Biology (June, 1973).
Kirchner, James W. “The Gaia Hypotheses: Are They Testable? Are They
Useful?” In Scientists on Gaia, edited by Stephen H. Schneider and
Penelope J. Boston. Cambridge, Mass.: MIT Press, 1991.
Milne, Lorus Johnson. The Balance of Nature. New York: Knopf, 1960.
31
BIODIVERSITY
Types of ecology: Community ecology; Ecosystem ecology; Global
ecology; Population ecology; Restoration and conservation ecology
The Wildlife Society has defined biodiversity as “the richness, abundance, and
variability of plant and animal species and communities and the ecological process
that link them with one another and with soil, air, and water.”
M any kinds of specialists—including organismic biologists, popu-

lation and evolutionary biologists, geneticists, and ecologists—
investigate biological processes that are encompassed by the concept of
biodiversity. Conservation biologists are concerned with the totality of bio-
diversity, including the process of speciation that forms new species, the
measurement of biodiversity, and factors involved in the extinction pro-
cess. However, the primary thrust of their efforts is the development of
strategies to preserve biodiversity. The biodiversity paradigm connects
classical taxonomic and morphological studies of organisms with modern
techniques employed by those working at the molecular level.
It is generally accepted that biodiversity can be approached at three lev-
els of organization, commonly identified as species diversity, ecosystem
diversity, and genetic diversity. Some also recognize biological phenomena
diversity.
Species Diversity
No one knows how many species inhabit the earth. Estimates range from
five million to several times that number. Each species consists of individu-
als that are somewhat similar and capable of interbreeding with other
members of their species but are not usually able to interbreed with indi-
viduals of other species. The species that occupy a particular ecosystem are
a subset of the species as a whole. Ecosystems are generally considered to
be local units of nature; ponds, forests, and prairies are common examples.
Conservation biologists measure the species diversity of a given ecosys-
tem by first conducting a careful, quantitative inventory. From such data,
scientists may determine the “richness” of the ecosystem, which is simply
a reflection of the number of species present. Thus, an island with three
hundred species would be 50 percent richer than another with only two
hundred species. Some ecosystems, especially tropical rain forests and
coral reefs, are much richer than others. Among the least rich are tundra re-
gions and deserts.
32
Biodiversity
A second aspect of species diversity is “evenness,” defined as the de-

gree to which each of the various elements are present in similar percent-
ages of the total species. As an example, consider two forests, each of
which has a total of twenty species of trees. Suppose that the first forest has
a few tree species represented by rather high percentages and the remain-
der by low percentages. The second forest, with its species more evenly
distributed, would rate higher on a evenness scale.
Species diversity, therefore, is a value that combines measures of both
species richness and species evenness. Values obtained from a diversity in-
dex are used in comparing species diversity among ecosystems of both the
same and different types. They also have implications for the preservation
of ecosystems; other things being equal, it would be preferable to preserve
ecosystems with a high diversity index, thus protecting a larger number of
species.
Species Diversity and Geographical Areas

Considerable effort has been expended to predict species diversity as de-
termined by the nature of the area involved. For example, island biogeog-
raphy theory suggests that islands that are larger, nearer to other islands or
continents, and have a more heterogeneous landscape would be expected
to have a higher species diversity than those possessing alternate traits.
Such predictions apply not only to literal islands but also to other discon-
tinuous ecosystems. Examples would be the ecosystems of isolated moun-
taintops in alpine tundra or those of ponds several miles apart.
The application of island biogeography theory to designing nature pre-
serves was proposed by Jared Diamond in 1975. His suggestion began the
“single larger or several smaller,” or SLOSS, area controversy. Although is-
land biogeography theory would, in many instances, suggest selecting one
large area for a nature preserve, it is often the case that several smaller ar-
eas, if carefully selected, could preserve more species.
The species diversity of a particular ecosystem is subject to change over
time. Pollution, deforestation, and other types of habitat degradation invari-
ably reduce diversity. Conversely, during the extended process of ecological
succession that follows disturbances, species diversity typically increases
until a permanent, climax ecosystem with a large index of diversity results.
It is generally assumed by ecologists that more diverse ecosystems are more
stable than are those with less diversity. Certainly, the more species pres-
ent, the greater the opportunity for various interactions, both with other
species and with the environment. Examples of interspecific reactions in-
clude mutualism, predation, and parasitism. Such interactions apparently
help to integrate a community into a whole, thus increasing its stability.
33
Biodiversity
Ecosystem Diversity
Ecology can be defined as the study of ecosystems. From a conservation
standpoint, ecosystems are important because they sustain their particular
assemblage of living species. Conservation biologists also consider ecosys-
tems to have an intrinsic value beyond the species they harbor. Therefore,
it would be ideal if representative global ecosystems could be preserved.
However, this is far from realization. Just deciding where to draw the
line between interfacing ecosystems can be a problem. For example, the
water level of a stream running through a forest is subject to seasonal fluc-
tuation, causing a transitional zone characterized by the biota from both
adjoining ecosystems. Such ubiquitous zones negate the view that ecosys-
tems are discrete units with easily recognized boundaries.
The protection of diverse ecosystems is of utmost importance to the
maintenance of biodiversity. However, ecosystems throughout the world
are threatened by global warming, air and water pollution, acid deposi-
tion, ozone depletion, and other destructive forces. At the local level, de-
forestation, thermal pollution, urbanization, and poor agricultural prac-
tices are among the problems affecting ecosystems and therefore reducing
biodiversity. Both global and local environmental problems are amplified
by rapidly increasing world population pressures.
In the process of determining which ecosystems are most in need of pro-
tection, it has become apparent to many scientists that a system for naming
and classifying ecosystems is highly desirable, if not imperative. Efforts
are being made to establish a system similar to the hierarchical system ap-
plied to species that was developed by Swedish botanist Carl Linnaeus
during the eighteenth century. However, a classification system for ecosys-
tems is far from complete. Freshwater, marine, and terrestrial ecosystems
are recognized as main categories, with each further divided into particu-
lar types. Though tentative, this has made possible the identification and
preservation of a wide range of representative, threatened ecosystems.
In 1995 conservation biologist Reed F. Noss of Oregon State University
and his colleagues identified more than 126 types of ecosystems in the
United States that are threatened or critically endangered. The following
list illustrates their diversity: southern Appalachian spruce-fir forests;
eastern grasslands, savannas, and barrens; California native grasslands;
Hawaiian dry forests; caves and Karst systems; old-growth forests of the
Pacific Northwest; and southern forested wetlands.
Not all ecosystems can be saved. Establishing priorities involves many
considerations, some of which are economic and political. Ideally, choices
would be made on merit: rarity, size, number of endangered species they
include, and other objective, scientific criteria.
34
Biodiversity
Genetic Diversity
Most of the variation among individuals of the same species is caused by
the different genotypes (combinations of genes) that they possess. Such ge-
netic diversity is readily apparent in cultivated or domesticated species
such as cats, dogs, and corn, but also exists, though usually to a lesser de-
gree, in wild species. Genetic diversity can be measured only by exacting
molecular laboratory procedures. The tests detect the amount of variation
in the deoxyribonucleic acid (DNA) or isoenzymes (chemically distinct en-
zymes) possessed by various individuals of the species in question.
A significant degree of genetic diversity within a population or species
confers a great advantage. This diversity is the raw material that allows
evolutionary processes to occur. When a local population becomes too
small, it is subject to a serious decline in vigor from increased inbreeding.
This leads, in turn, to a downward, self-perpetuating spiral in genetic di-
versity and further reduction in population size. Extinction may be immi-
nent. In the grand scheme of nature, this is a catastrophic event; never
again will that particular genome (set of genes) exist anywhere on the
earth. Extinction is the process by which global biodiversity is reduced.
Biological Phenomena Diversity

Biological phenomena diversity refers to the numerous unique biological
events that occur in natural areas throughout the world. Examples include
the congregation of thousands of monarch butterflies on tree limbs at Point
Pelee in Ontario, Canada, as they await favorable conditions before contin-
uing their migration, or the return of hundreds of loggerhead sea turtles
each April to Padre Island in the Gulf of Mexico in order to lay their eggs.
Conservation Biology
Although biologists have been concerned with protecting plant and ani-
mal species for decades, only recently has conservation biology emerged
as an identifiable discipline. Conceived in a perceived crisis of biological
extinctions, conservation biology differs from related disciplines, such as
ecology, because of its advocative nature and its insistence on maintaining
biodiversity as intrinsically good. Conservation biology is a value-laden
science, and some critics consider it akin to a religion with an accepted
dogma.
The prospect of preserving global ecosystems and the life processes
they make possible, all necessary for maintaining global diversity, is not
promising. Western culture does not give environmental concerns a high
priority. For those who do, there is more often a concern over issues relat-
ing to immediate health effects than concern over the loss of biodiversity.
35
Biodiversity
Only when education in basic biology and ecology at all levels is extended
to include an awareness of the importance of biodiversity will there de-
velop the necessary impetus to save ecosystems and all their inhabitants,
including humans.
Thomas E. Hemmerly
See also: Animal-plant interactions; Biogeography; Biomes: determinants;

Biomes: types; Biosphere concept; Communities: ecosystem interactions;
Communities: structure; Conservation biology; Ecosystems: definition
and history; Ecosystems: studies; Endangered animal species; Endangered
plant species; Extinctions and evolutionary explosions; Gene flow; Genetic
diversity; Habitats and biomes; Restoration ecology; Species loss; Zoos.

Baskin, Y. “Ecologists Dare to Ask: How Much Does Diversity Matter?”
Science 264 (April 8, 1994): 202-203.
Burton, John, ed. The Atlas of Endangered Species. 2d ed. New York:
Macmillan, 1999.
Cracraft, Joel, and Francesca T. Grifo, eds. The Living Planet in Crisis: Biodi-
versity Science and Policy. Foreword by Edward O. Wilson. New York:
Columbia University Press, 1999.
DiSilvestro, Roger L. The Endangered Kingdom: The Struggle to Save America’s
Wildlife. New York: John Wiley & Sons, 1989.
Ehrlich, Paul, and Anne Ehrlich. Extinction: The Causes and Consequences of
the Disappearance of Species. New York: Random House, 1981.
National Research Council. Science and the Endangered Species Act. Wash-
ington, D.C.: National Academy Press, 1995.
New, T. R. An Introduction to Invertebrate Conservation. New York: Oxford
Raven, Peter H., and George B. Johnson. Biology. 4th ed. Boston: McGraw-
Hill, 1996.
Reaka-Kudla, Marjorie L., Don E. Wilson, and Edward O. Wilson, eds. Bio-
diversity II: Understanding and Protecting Our Biological Resources. Wash-
ington, D.C.: Joseph Henry Press, 1997.
Ricketts, Taylor H., et al. Terrestrial Ecoregions of North America: A Conserva-
tion Assessment. Washington, D.C.: Island Press, 1999.
Stefoff, Rebecca. Extinction. New York: Chelsea House, 1992.
Tudge, Colin. The Variety of Life: The Meaning of Biodiversity. New York: Ox-
ford University Press, 2000.
Wilson, Edward O., ed. Biodiversity. Washington, D.C.: National Academy
Press, 1988.
36
BIOGEOGRAPHY
Types of ecology: Community ecology; Population ecology
To understand the underlying geography of plant and animal distributions,

biogeographers integrate considerations of historical and current events and con-
ditions.
B iogeography is the science that seeks to understand spatial patterns of

biodiversity. By examining past and present distributions, biogeog-
raphers attempt to explain why certain groups of organisms occur where
they do, what enables them to live there, -and what factors prevent them
from moving or living elsewhere. To address these issues, biogeographic
investigations must consider the effects of climate, topography, and other
kinds of organisms, as well as historical events such as tectonic effects and
glaciation. Such historical considerations are frequently important and
conditions are constantly changing, so a group’s closest relatives and their
distribution must be taken into account in order to accommodate evolu-
tionary events.
Because biogeography is such a broad discipline, individuals can rarely
address an entire spectrum of relevant questions. Consequently, most
biogeographers become specialists. For example, phytogeographers study
plant distributions and zoogeographers examine those of animals. Histori-
cal biogeographers reconstruct origins, dispersal events, and extinctions
through time. Ecological biogeographers concentrate on interactions be-
tween organisms and their environments to explain distribution patterns,
and paleoecologists try to bridge the gap between historical and current
conditions. Also related to the breadth of the discipline is the necessity for
biogeographers to be conversant in one or more related fields. A broad
knowledge of biology is obviously fundamental, as is an understanding of
physical geography, but geology, paleontology, and climatology, among
others, may be equally important.
Trends in Biogeography
Several consistent trends have emerged from biogeographic studies. Com-
munities in isolated regions (especially large islands that have not been in
contact with continents for long periods of time) tend to be unlike those
found anywhere else. More types of organisms are found in tropical than
in temperate or arctic regions. Fewer types of organisms are found on oce-
anic islands, although the organisms that are present may be found in phe-
37
Biogeography
nomenal densities. On the other hand, some inconsistencies are striking.

For example, some groups of related organisms are found throughout the
world whereas other groups have very restricted ranges.
Generally speaking, groups of organisms that are broadly distributed
are either very old (their ancestors were in place before the continents
drifted apart) or very mobile. Mobile organisms may be able to disperse ac-
tively, that is, on their own power (some species of birds and large marine
mammals are good examples), whereas others are dispersed passively.
Seeds of many plants, microscopic planktonic organisms, and insects are
often transported by wind, currents, or other organisms. Limits to the dis-
persal of organisms are myriad. Size limits the ability to be blown by the
wind; for example, dandelion thistles are more readily dispersed by even
mild breezes than walnuts. If water is a factor, buoyancy is critical. Coco-
nuts are found on tropical shorelines around the world in large part be-
cause they do not sink, nor are their hard shells easily penetrated by salt
water. Physical barriers often limit dispersal. Deserts effectively block or-
ganisms that require moisture, land halts movement of aquatic forms, and
mountains prevent the passage of plants and animals that cannot tolerate
the conditions associated with high elevations. Some barriers even appear
to be “psychological”; birds that could easily fly cross a stream or lake, for
example, often will not.
Island Biogeography
In 1967, Robert H. MacArthur and Edward O. Wilson published a classic
volume titled The Theory of Island Biogeography. Although islands have fig-
ured prominently in modern biology (Charles Darwin and Alfred Russel
Wallace both relied heavily on evidence from islands when formulating
their theories of natural selection), only during the last third of the twenti-
eth century was island biogeography recognized as a distinct discipline.
Many of the principles that form the foundation of biogeography ema-
nated from studies of islands. Among these are relationships between bio-
diversity and island size, ecological heterogeneity, and proximity to conti-
nents; between isolation and endemism (species evolving in a given area
and found nowhere else in the world); and between island size and loca-
tion and rates of immigration, colonization, and extinction. In addition, is-
land biogeographers frequently have been at the center of debates arguing
the relevance of dispersal versus vicariance. Central to these disputes is
whether disjunct distributions of organisms are attributable to movement
over barriers (dispersal) or to the creation of a barrier that separated a pre-
viously contiguous range (vicariance). Although both have undoubtedly
played important roles, the debate rages over which was primarily respon-
38
Biogeography
sible for the current distributions of many faunas, both on islands sur-
rounded by water or on terrestrial “islands” surrounded by other inhospi-
table habitats.
Robert Powell
See also: Adaptations and their mechanisms; Adaptive radiation; Biodi-

versity; Biological invasions; Clines, hybrid zones, and introgression;
Communities: ecosystem interactions; Communities: structure; Evolution:
definition and theories; Food chains and webs; Gene flow; Genetic diver-
sity; Genetic drift; Isolating mechanisms; Landscape ecology; Migration;
Nonrandom mating, genetic drift, and mutation; Population genetics; Spe-
ciation; Species loss.

Cox, C. Barry, and Peter D. Moore. Biogeography: An Ecological and Evolu-
tionary Approach. 6th ed. Malden, Mass.: Blackwell Science, 2000.
Groombridge, Brian, and M. K. Jenkins, eds. Global Biodiversity: Earth’s Liv-
ing Resources in the Twenty-first Century. Cambridge, England: World
Conservation Monitoring Centre, 2000.
Myers, Alan A., and Paul S. Giller, eds. Analytical Biogeography: An Inte-
grated Approach to the Study of Animal and Plant Distributions. New York:
Chapman & Hall, 1988.
Whittaker, Robert J. Island Biogeography: Ecology, Evolution, and Conserva-
tion. New York: Oxford University Press, 1998.
Wilson, Edward O., ed. Biodiversity. Washington, D.C.: National Academic
Press, 1988.
39
BIOLOGICAL INVASIONS
Types of ecology: Community ecology; Ecosystem ecology;
Ecotoxicology
Biological invasions are the entry of a type of organism into an ecosystem outside
its historic range. In a biological invasion, the “invading” organism may be an in-
fectious virus, a bacterium, a plant, an insect, or an animal.
S pecies introduced to an area from somewhere else are referred to as

alien or exotic species or as invaders. Because an exotic species is not
native to the new area, it is often unsuccessful in establishing a viable pop-
ulation and disappears. The fossil record, as well as historical documenta-
tion, indicates that this is the fate of many species in new environments as
they move from their native habitats. Occasionally, however, an invading
species finds the new environment to its liking. In this case, the invader
may become so successful in exploiting its new habitat that it can com-
pletely alter the ecological balance of an ecosystem, decreasing biodiver-
sity and altering the local biological hierarchy. Because of this ability to al-
ter ecosystems, exotic invaders are considered major agents in driving
native species to extinction.
Biological invasions by notorious species constitute a significant com-
ponent of earth’s history. In general, large-scale climatic changes and geo-
logical crises are at the origin of massive exchanges of flora and fauna. On a
geologic time scale, migrations of invading species from one continent to
another are true evolutionary processes, just as speciation and extinction
are. On a smaller scale, physical barriers such as oceans, mountains, and
deserts can be overcome by many organisms as their populations expand.
Organisms can be carried by water in rivers or ocean currents, transported
by wind, or carried by other species as they migrate seasonally or to escape
environmental pressures. Humans have transplanted plants since the be-
ginning of plant cultivation in pre-Columbian times. The geological and
historical records of the earth suggest that biological invasions contribute
substantially to an increase in the rate of extinction within ecosystems.
Invasive Plants
In modern times, most people are not aware of the distinction between na-
tive plants and exotic species growing in their region. Recent increases in
intercontinental invasion rates by exotic species, brought about primarily
by human activity, create important ecological problems for the recipient
40
Biological invasions
lands. Invasive plants in North America include eucalyptus trees, morning

glory, and pampas grass.
It would seem logical to assume that invading species might add to the
biodiversity of a region, but many invaders have the opposite effect. In all
ecosystems the new species are often opportunistic, driving out native spe-
cies by competing with them for resources. For example, Pueraria lobata, or
kudzu, is a vine native to Japan. Introduced in the United States at the 1876
Philadelphia Exposition, kudzu was planted to control erosion on hillsides
and for livestock forage. By the end of the twentieth century, it could be
found from Connecticut to Missouri, extending south to Texas and Florida.
Kudzu covers everything in its path and grows as much as 1 foot (0.3 me-
ter) per day. Similarly, English ivy (Hedera helix), a native of Eurasia, is con-
sidered a serious problem in West Coast states. It forms “ivy deserts” in
forests and crowds out native trees and shrubs that make up essential
wildlife habitat.
The invasion of an ecosystem by an exotic species can effectively alter
ecosystem processes. An invading species does not simply consume or
compete with native species but can actually change the rules of existence
within the ecosystem by altering processes such as primary productivity,
decomposition, hydrology, geomorphology, nutrient cycling, and natural
disturbance regimes.
Invasive Insects and Microorganisms

The invasion of native forests alone by nonnative insects and microorgan-
isms has been devastating on many continents. The white pine blister rust
and the balsam woolly adelgid have invaded both commercial and pre-
served forest lands in North America. Both exotics were brought to North
America in the late 1800’s on nursery stock from Europe. The balsam
woolly adelgid attacks fir trees and causes their death within two to seven
years from chemical damage and by feeding on the tree’s vascular tissue.
The adelgid has killed nearly every adult cone-bearing fir tree in the south-
ern Appalachian Mountains. The white pine blister rust attacks five-needle
pines; in the western United States fewer than 10 pine trees in 100,000 are
resistant. Because white pine seeds are an essential food source for bears
and other animals, the loss of the trees is having severe consequences
across the food chain.
Since the 1800’s the deciduous trees of eastern North America have
been attacked numerous times by waves of invading exotic species and
diseases. One of the most notable invaders is the gypsy moth, which con-
sumes a variety of tree species. Other invaders have virtually eliminated
the once-dominant American chestnut and the American elm. Tree species
41
Biological invasions
that continue to decline because of new invaders include the American

beech, mountain ash, white birch, butternut, sugar maple, flowering dog-
wood, and eastern hemlock. It is widely accepted that the invasion of ex-
otic species is the single greatest threat to the diversity of deciduous forests
in North America.
Effects on Humans and Humans as Invaders

Some introduced exotic species are beneficial to humanity. It would be im-
possible to support the present world human population entirely on spe-
cies native to their regions. Humans, the ultimate biological invaders, have
been responsible for the extinction of many species and will continue to be
in the future. At the beginning of the twenty-first century, the United States
was spending $4 billion annually to eradicate invasive plant species, a fig-
ure that does not take into account loss of biodiversity or wildlife habitat.
Randall L. Milstein, updated by Elizabeth Slocum
See also: Biodiversity; Biogeography; Biomagnification; Communities:

ecosystem interactions; Deforestation; Endangered animal species; Endan-
gered plant species; Eutrophication; Invasive plants; Succession.

Bright, Chris. Life out of Bounds: Bioinvasion in a Borderless World. New York:
Norton, 1998.
Cox, George W. Alien Species in North America and Hawaii: Impacts on Natural
Ecosystems. Washington, D.C.: Island Press, 1999.
Crosby, Alfred. Ecological Imperialism: The Biological Expansion of Europe,
900-1900. New York: Cambridge University Press, 1994.
Drake, J. A., et al. Biological Invasions: A Global Perspective. New York: Wiley,
1989.
Hengeveld, Rob. Dynamics of Biological Invasions. New York: Chapman and
Hall, 1989.
42
BIOLUMINESCENCE
Types of ecology: Chemical ecology; Physiological ecology
Bioluminescence is visible light emitted from living organisms. Half of the orders
of animals include luminescent species. These organisms are widespread, occur-
ring in marine, terrestrial, and freshwater habitats. Bioluminescence is used for
defense, predation, and communication.
B ioluminescence is the visible light produced by luminous animals,

plants, fungi, protists, and bacteria that results from a biochemical re-
action with oxygen. Unlike incandescent light from electric light bulbs,
bioluminescence is produced without accompanying heat. Biolumines-
cence was first described in 500 b.c.e., but the chemical mechanism of bio-
luminescence was not elucidated until the beginning of the twentieth cen-
tury. The ability to luminesce appears to have arisen as many as thirty
times during evolution. The chemical systems used by luminescent organ-
isms are similar but not exactly the same. Most organisms use a luciferin/
luciferase system. The luciferin molecules are oxidized through catalysis
by an oxidase enzyme (luciferase). The oxidized form of luciferin is in an
excited electronic state that relaxes to the ground state through light emis-
sion.
Types of Bioluminescence
Animals may produce light in one of three ways. The bioluminescence
may be intracellular: Chemical reactions within specialized cells result in
the emission of visible light. These specialized cells are often found within
photophores. These light-producing organs may be arranged in symmetri-
cal rows along the animal’s body, in a single unit overhanging the mouth,
or in patches under the eyes, and are connected to the nervous system. Al-
ternatively, the bioluminescence may be extracellular: The animals secrete
chemicals that react in their surroundings to produce light. The third op-
tion involves a symbiotic relationship between an animal and biolumi-
nescent bacteria. Several species of fish and squid harbor bioluminescent
bacteria in specialized light organs. The symbiotic relationship is specific:
Each type of fish or squid associates with a certain type of bacteria. The
bacteria-filled organ is continuously luminous. The animal regulates the
emission of light either by melanophores scattered over the surface of
the organ or by a black membrane that may be mechanically drawn over
the organ.
43
Bioluminescence
Image Not Available
Although bioluminescence is widespread in animals, its occurrence is

sporadic. Most of the bioluminescent animal species are invertebrates.
Among the vertebrates, only fish exhibit bioluminescence; there are no
known luminous amphibians, reptiles, birds, or mammals. Although bio-
luminescence is found in terrestrial and freshwater environments, the ma-
jority of luminous organisms are marine. Scientists estimate that 96 percent
of all creatures in the deep sea possess some form of light generation.
Functions of Bioluminescence
There appear to be three main uses of bioluminescence: finding or attract-
ing prey, defense against predators, and communication. Although visible
light penetrates into the ocean to one thousand meters at most, most fish
living below one thousand meters possess eyes or other photoreceptors.
Many deep-sea fish have dangling luminous light organs to attract prey.
Terrestrial flies have also exploited bioluminescence for predation. The
glow of glowworms (fly larvae) living in caves serves to attract insect prey,
which get snared in the glowworms’ sticky mucous threads. Fungus gnats
(carnivorous flies) attract small arthropods through light emission and
capture the prey in webs of mucus and silk.
44
Bioluminescence
Bioluminescence can serve as a decoy or camouflage. For example, jel-

lyfish, such as comb jellies, produce bright flashes to startle a predator,
while siphonophores can release thousands of glowing particles into the
water as a mimic of small plankton to confuse the predator. Other jellyfish
produce a glowing slime that can stick to a potential predator and make it
vulnerable to its predators. Many squid and some fish possess photo-
phores that project light downward, regardless of the orientation of the an-
imal’s body. The emitted light matches that of ambient light when viewed
from below, rendering the squid invisible to both predators and prey.
The best-known example of bioluminescence used as communication is
in fireflies, the common name for any of a large family of luminescent bee-
tles. Luminescent glands are located on the undersides of the rear abdomi-
nal segments. There is an exchange of flashes between males and females.
Females respond to the flashes of flying males, with the result that the male
eventually approaches the female for the purpose of mating. To avoid con-
fusion between members of different types of fireflies, the signals of each
species are coded in a unique temporal sequence of flashing, the timing of
which is controlled by the abundant nerves in the insect’s light-making or-
gan. Females of one genus of fireflies (Photuris) take advantage of this by
mimicking the response of females of another genus (Photinus) to lure
Photinus males that the Photuris females then kill and eat.
Some marine animals also utilize bioluminescence for communication.
For example, lantern fish and hatchetfish (the most abundant vertebrate on
earth) possess distinct arrangements of light organs on their bodies that
can serve as species- and sex-recognition patterns; female fire worms re-
lease luminescent chemicals into the water during mating, beginning one
hour after sundown on the three nights following the full moon; and deep-
sea dragonfish emit red light that is undetectable except by other drag-
onfish.
Lisa M. Sardinia
See also: Adaptations and their mechanisms; Camouflage; Communica-

tion; Defense mechanisms; Marine biomes; Metabolites; Mimicry; Phero-
mones; Poisonous animals; Poisonous plants; Pollination; Predation; Tro-
pisms.

Presnall, Judith Janda. Animals That Glow. Salem, Mass.: Franklin Watts,
1993.
Robison, Bruce H. “Light in the Ocean’s Midwaters: Bioluminescent Ma-
rine Animals.” Scientific American 273 (July, 1995): 60-64.
45
Bioluminescence
Silverstein, Alvin, and Virginia Silverstein. Nature’s Living Lights: Fireflies

and Other Bioluminescent Creatures. Boston: Little, Brown, 1988.
Toner, Mike. “When Squid Shine and Mushrooms Glow, Fish Twinkle, and
Worms Turn into Stars.” International Wildlife 24 (May/June, 1994): 30-
37.
Tweit, Susan J. “Dance of the Fireflies.” Audubon 101 (July/August, 1999):
26-30.
46
BIOMAGNIFICATION
Type of ecology: Ecotoxicology
Biomagnification is the accumulation of toxic contaminants in the environment

as they move up through the food chain. As members of each level of the food chain
are progressively eaten by those organisms found in higher levels of the chain, the
concentration of toxic chemicals within the tissues of the higher organisms in-
creases.
N ot all chemicals, potentially toxic or not, are equally likely to undergo

biomagnification. However, molecules susceptible to biomagnifi-
cation have certain characteristics in common. They are resistant to natural
microbial degradation and therefore persist in the environment. They are
also lipophilic, tending to accumulate in the fatty tissue of organisms. In
addition, the chemical must be biologically active in order to have an effect
on the organism in which it is found. Such compounds are likely to be ab-
sorbed from food or water in the environment and stored within the mem-
branes or fatty tissues.
Pesticides
The process usually begins with the spraying of pesticides for the purpose
of controlling insect populations. Industrial contamination, including the
release of heavy metals, can be an additional cause of such pollution. Bio-
magnification results when these chemicals contaminate the water supply
and are absorbed into the lipid membranes of microbial organisms. This
process, often referred to as bioaccumulation, results in the initial concen-
tration of the chemical in an organism in a form that is not naturally ex-
creted with normal waste material. Levels of the chemical may reach any-
where from one to three times that found in the surrounding environment.
Since the nature of the chemical is such that it is neither degraded nor ex-
creted, it remains within the organism.
As organisms on the bottom of the food chain are eaten and digested by
members of the next level in the chain, the concentration of the accumu-
lated material significantly increases; at each subsequent level, the concen-
tration may reach one order of magnitude (a tenfold increase) higher. Con-
sequently, the levels of the pollutant at the top of the environmental food
chain for the ecosystem in question—such as fish, carnivorous birds, or hu-
mans—may be as much as one million times more concentrated than the
original, presumably safe, levels in the environment.
47
Biomagnification
DDT
For example, studies of dichloro-diphenyl-trichloroethane (DDT) levels in
the 1960’s found that zooplankton at the bottom of the food chain had ac-
cumulated nearly one thousand times the level of the pollutant in the sur-
rounding water. Ingestion of the plankton by fish resulted in concentration
by another factor of several hundred. By the time the fish were eaten by
predatory birds, the level of DDT was concentrated by a factor of more
than two hundred thousand. DDT is characteristic of most pollutants sub-
ject to potential biomagnification. It is relatively stable in the environment,
persisting for decades. It is soluble in lipids and readily incorporated into
the membranes of organisms.
Since pesticides are, by their nature, biologically active compounds,
which reflects their ability to control insects, they are of particular con-
cern if subject to biomagnification. DDT remains the classical example
of how bioaccumulation and biomagnification may have an effect on the
environment. Initially introduced as a pesticide for control of insects and
insect-borne disease, DDT was not thought to be particularly toxic. How-
ever, biomagnification of the chemical was found to result in the deaths
of birds and other wildlife. In addition, DDT contamination was found
to result in formation of thin egg shells that greatly reduced the birth-
rate among birds. Before the use of DDT was banned in the 1960’s, the
population levels of predatory birds such as eagles and falcons had
fallen to a fraction of the levels found prior to use of the insecticide.
Though it was unclear whether there was any direct effect on the hu-
man population in the United States, the discovery of elevated levels of
DDT in human tissue contributed to the decision to ban the use of the
chemical.
Other Toxic Pesticides

While DDT represents the classic example of biomagnification of a toxic
chemical, it is by no means the only representative of potential environ-
mental pollutants. Other pesticides with similar characteristics include
pesticides such as aldrin, chlordane, parathion, and toxaphene. In addi-
tion, cyanide, polychlorinated biphenyls (PCBs), and heavy metals—such
as selenium, mercury, copper, lead, and zinc—have also been found to con-
centrate within the food chain.
Some heavy metals are inherently toxic or may undergo microbial mod-
ification to increase their toxic potential. For example, mercury does not
naturally accumulate in membranes and was therefore not originally
viewed as a significant danger to the environment. However, some micro-
organisms are capable of adding a methyl group to the metal and produc-
48
Biomagnification
ing methyl mercury, a highly toxic material that does accumulate in fatty
tissue and membranes.
Prevention
Several procedures have been adopted since the 1960’s to prevent the bio-
magnification of toxic materials. In addition to outright bans, pesticides
are often modified to prevent their accumulation in the environment. Most
synthetic pesticides contain chemical structures that are easily degraded
by microorganisms found in the environment. Ideally, the pesticide should
survive no longer than a single growing season before being rendered
harmless by the environmental flora. Often such chemical changes require
only simple modification of the basic structure.
Richard Adler
See also: Acid deposition; Biological invasions; Biopesticides; Endangered

animal species; Food chains and webs; Genetically modified foods; Inte-
grated pest management; Ocean pollution and oil spills; Pesticides;
Phytoplankton; Pollution effects; Waste management.

Atlas, Ronald, and Richard Bartha. Microbial Ecology: Fundamentals and Ap-
plications. Redwood City, Calif.: Benjamin/Cummings, 1993.
Carson, Rachel. Silent Spring. Boston: Houghton Mifflin, 1962.
Colborn, Theo, Dianne Dumanoski, and John Peterson Myers. Our Stolen
Future: Are We Threatening Our Fertility, Intelligence, and Survival? A Sci-
entific Detective Story. New York: Dutton, 1996.
49
BIOMASS RELATED TO ENERGY
Type of ecology: Ecoenergetics
The relationship between the accumulation of living matter resulting from the pri-
mary production of plants or the secondary production of animals (biomass) and
the energy potentially available to other organisms in an ecosystem forms the basis
of the study of biomass related to energy.
B iomass is the amount of organic matter, such as animal and plant tis-
sue, found at a particular time and place. The rate of accumulation of
biomass is termed productivity. Primary production is the rate at which
plants produce new organic matter through photosynthesis. Secondary
production is the rate at which animals produce their organic matter by
feeding on other organisms. Biomass is an instantaneous measure of the
amount of organic matter, while primary and secondary production give
measures of the rates at which biomass increases. Plant and animal bio-
mass consists mostly of carbon-rich molecules, such as sugars, starches,
proteins, and lipids, and other substances, such as minerals, bone, and
shell. The carbon-rich organic molecules are not only the building blocks of
life but also the energy-rich molecules used by organisms to fuel their ac-
tivities.
Primary Production: Photosynthesis

Ultimately, all energy used by organisms to produce the building blocks of
life and to drive life processes originated as solar energy captured by
plants. Only a small fraction, less than 2 percent, of the total solar light en-
ergy received by a plant is absorbed and transformed by photosynthesis
into energy-containing organic molecules. The rest of the sun’s energy
passes out of the plant as heat. The rate at which plants capture light en-
ergy and transform it into chemical energy is called primary production.
Because plants do not rely on other organisms to provide their energy
needs, they are referred to as primary producers, or autotrophs (meaning
“self-feeding”). In addition to light energy, plants must absorb water, car-
bon dioxide gas, and simple nutrients, such as nitrate and phosphate, to
produce various organic molecules during photosynthesis. Oxygen gas is
also produced.
Sugars are the first energy-containing organic molecules produced in
photosynthesis, and they can be changed to other, more complex, mole-
cules, such as starches, proteins, and fats. The energy in the sugar mole-
50
Biomass related to energy
cules can be used immediately by the plants to maintain their own respira-
tion needs, stored as starches and fats, or can be converted to new plant
tissue. It is the stored organic matter plus new tissue that contributes to the
growth of plants and to biomass.
Because the energy-containing products of photosynthesis can be used
either immediately in respiration or in the formation of new plant biomass,
two types of primary production can be distinguished. Gross production
refers to the total amount of energy produced by photosynthesis. It in-
cludes both the energy used by the plant for respiration and the energy that
goes into new biomass. Net production refers only to the amount of energy
that accumulates as new biomass. It is only the energy in net production
that is potentially available to animal consumers as food.
The rate of primary production varies directly with the rate of photo-
synthesis; therefore, factors in the environment that affect the rate of pho-
tosynthesis affect the rate of primary production. These factors most often
Transformation of Sunlight
into Biochemical Energy
Light energy
from the sun
Chloroplasts in
plant cells convert
light, water, and carbon
dioxide into carbohydrates
via photosynthesis
Oxygen, as a
by-product of
photosynthesis,
is released
Respiration by-products
are carbon dioxide,
water, and energy
dissipated as heat
Mitochondria in plant cells

perform cellular respiration
Respiration results in energy

storage in ATP molecules
51
include light intensity, temperature, nutrient concentrations, and moisture

conditions. Each species of plant has a specific combination of these factors
that promotes maximum rates of primary production. If one or more of
these factors is in excess or is in short supply, then the rate of primary pro-
duction is slowed.
On land, the rate of primary production by plants is determined largely
by light, temperature, and rainfall. The favorable combination of intense
sunlight for twelve hours per day, warm temperatures throughout the
year, and considerable rainfall make the tropical rain forests the most pro-
ductive ecosystem on land. In contrast, Arctic tundra vegetation is ex-
posed to reduced light intensity, very cold winters, and cool summers. Pri-
mary production there is very low. In deserts, the lack of water severely
limits primary production even though light and temperature are other-
wise favorable.
In aquatic habitats, rates of primary production by algae, such as
phytoplankton, are determined by nutrient concentration and light inten-
sity. As sunlight penetrates water, it is quickly absorbed by the water mole-
cules and by small suspended particles. Thus, all primary production oc-
curs near the surface, as long as nutrients are available. Although the
waters of the open ocean are very clear, and sunlight can penetrate to great
depths, the scarcity of nutrients reduces the rate of primary production to
less than one-tenth that of coastal bays.
Secondary Production
The energy and material needs of some organisms are met by consuming
the organic materials produced by others. These consumer organisms are
called heterotrophs; there are two types. Those that obtain their food from
other living organisms are called consumers and include all animals.
Those that obtain their energy from dead organisms are called decom-
posers and include mostly the fungi and bacteria.
The energy available to each type of consumer becomes progressively
less at each level of the food chain. Each consumer level uses most of
its food energy, about 90 percent, to fuel its respiratory activities. In this
energy-releasing process, most of the food energy is actually converted to
heat and is lost to the environment. Only 10 percent or less of the original
food energy is used to form new biomass. It is only this small amount of en-
ergy that is available for the next consumer level. The result is that food
chains are limited in their number of links or levels by the reduced amount
of energy available at each higher level.
Generally, the greater the amount of primary production, the larger the
number of consumer organisms and the longer the food chain. Most food
52
chains consist of three levels; rarely are there examples of up to five levels.
It should be noted that the food chain concept is a simplified view of a
more complex network of energy pathways, known as food webs, that oc-
cur in nature. Another outcome of the reduction in energy flow up the food
chain is a progressive decrease in production and biomass. The most pro-
ductive level, and the one with the greatest biomass, is therefore the pri-
mary producers, or plants.
Human Threats to Primary Production

The total natural primary production of the earth is limited, and human ef-
forts to increase total world primary production much beyond its present
levels may be futile. One reason for this is that much of the earth’s surface
lacks optimal conditions for plant growth. The open ocean, which covers
about 71 percent of the earth’s surface, has very little plant growth. On
land, the Arctic, subarctic, and Antarctic regions are very unproductive
most of the year. Human attempts to increase primary production in the
form of food or fuel crops usually involve changing the characteristics of
the land, converting forests into croplands, for example, and adding large
quantities of nutrients and water. It has been estimated that humans are
currently utilizing most of the easily workable croplands and that the de-
velopment of additional lands for agriculture would require major
changes to currently unworkable habitats, changes that would be expen-
sive and demand much fuel energy.
The study of production processes is vitally important in understand-
ing the ecology of natural ecosystems. Such information is necessary to
manage and conserve habitats and their organisms in the face of human
pressures. These processes provide insight into the general health of eco-
systems. Pollutants, such as acid rain or industrial toxic wastes, are known
to reduce the primary and secondary productivity of forests and lakes.
Throughout the world, humans are reducing the biomass of the world’s
primary producers through deforestation. This is particularly true in the
tropics, where high population pressures have necessitated that land be
cleared for agriculture and development. There is a worldwide demand
for lumber. One obvious consequence is the dramatic reduction in the pri-
mary and secondary production of these areas. The clear-cutting (removal
of all the trees) of tropical forests allows unprotected soils to wash away
quickly during the heavy tropical rains. It will take hundreds of years for
new soils to develop and for the forest to return—if it can return at all.
Deforestation is also harmful in that tropical forests form a major part of
the world’s life-support system. For millions of years these forests have
buffered the earth’s atmosphere by producing the oxygen gas needed by
53
animals and by removing carbon dioxide and other toxic gases. The low
level of carbon dioxide in the atmosphere is believed to have moderated
the earth’s temperature, counteracting the so-called greenhouse effect. It is
therefore of great importance to understand and preserve these forests and
other primary producers of the world.
Ray P. Gerber
See also: Balance of nature; Communities: structure; Deforestation; Ecol-

ogy: definition; Ecosystems: studies; Food chains and webs; Geochemical
cycles; Herbivores; Hydrologic cycle; Nutrient cycles; Omnivores;
Phytoplankton; Predation; Rain forests and the atmosphere; Trophic levels

Brower, James E., and Jerrold H. Zar. Field and Laboratory Methods for Gen-
eral Ecology. 4th ed. Boston, Mass.: WCB McGraw-Hill, 1998.
Nybakken, James W. Marine Biology: An Ecological Approach. 5th ed. Menlo
Park, Calif.: Benjamin Cummings, 2001.
Odum, Howard T. Ecological and General Systems: An Introduction to Systems
Ecology. Rev. ed. Niwot: University Press of Colorado, 1994.
Pasztor, Janos, and Lars A. Kristoferson, eds. Bioenergy and the Environment.
Boulder, Colo.: Westview Press, 1990.
Ricklefs, Robert E. Ecology. 4th ed. New York: W. H. Freeman, 1999.
Smith, Robert L. Ecology and Field Biology. 5th ed. Menlo Park, Calif.: Addi-
son Wesley Longman, 1996.
54
BIOMES: DETERMINANTS
Types of ecology: Biomes; Ecosystem ecology; Global ecology;
Theoretical ecology
Biomes are large-scale ecological communities of both plants and animals, deter-
mined primarily by geography and climate. Worldwide, there are six major types
of biomes on land: forest, grassland, woodland, shrubland, semidesert scrub, and
desert.
O ne who travels latitudinally from the equator to the Arctic will cross
tropical forests, deserts, grasslands, temperate forests, coniferous for-
est, tundra, and ice fields. Those major types of natural vegetation at re-
gional scales are called biomes. A biome occurs wherever a particular set of
climatic and edaphic (soil-related) conditions prevail with similar physi-
ognomy. For example, prairies and other grasslands in the North Ameri-
can Middle West and West form a biome of temperate grasslands, where
moderately dry climate prevails. Tropical rain forests in the humid tropical
areas of South and Central America, Africa, and Southeast Asia create a
biome where rainfall is abundant and well-distributed through the year.
In general, biomes are delineated by both physiognomy and environ-
ment. There are six major physiognomic types on land: forest, grassland,
woodland, shrubland, semidesert scrub, and desert. Each of the six types
occurs in a wide range of environments. Therefore, more than one biome
may be defined within each physiognomic type according to major differ-
ences in climate. Tropical forests, temperate deciduous forest, and conifer-
ous forests are, for example, separate biomes, although forests dominate
all of them. On the other hand, some biome types, such as the tundra, are
dominated by a range of physiognomic types and are in one prevailing en-
vironmental region.
Classification of Biomes
There are many ways to classify biomes. One system, which designates a
small number of broadly defined biomes, divides global vegetation into
nine major terrestrial biomes: tundra, taiga, temperate forest, temperate
rain forest, tropical rain forest, savanna, temperate grasslands, chaparral,
and desert. Other systems more narrowly define biomes, designating a
larger total number. In those cases, some of the broadly defined biomes are
divided into two or more biomes. For example, the biome called temperate
forest in a broad classification may be separated into temperature decidu-
55
Biomes: determinants
Biomes and Their Features
1 2
Biome Annual Mean Rainfall Climate and Temperature
Desert 250 mm or less Arid, with extremes of heat

and cold
Grasslands 250-750 mm Cold winters, warm

summers; dry periods
Mediterranean Low to moderate Cool winters, hot summers;
scrub latitudes 30° to 45°; includes
chaparral, maquis
Rain forest (tropical) 2,500-4,500 mm 20-30°
Savanna, deciduous 1,500-2,500 mm Hot summers; 3-6 months
tropics dry; seasonal fires
Taiga (boreal forest) 1,000 mm Cold, long winters; mild,

short summers; seasonal
fires
Tundra Very low year-round Very cold (3° or less); soil
characerized by permafrost;
Arctic tundra occurs in
Arctic Circle; alpine tundra
in other high elevations
1
In millimeters
2
Degrees Celsius
ous forest and temperate evergreen forest in a fine classification. The bio-
me of desert in the broad classification may be broken into warm semi-
desert, cool semidesert, Arctic-alpine semidesert, Arctic-pine desert, and
true desert in the fine classification.
Description of Biome Distributions

Naturalists, geographers, and ecologists have tried to correlate world ma-
jor types of biomes to climatic patterns in both descriptive and quantitative
approaches. For example, in northern North America, the tundra and bo-
real forests are two broad belts of vegetation that stretch from east to west.
The distribution of the two biomes is primarily influenced by temperature.
South of those two belts are biome types that are mostly controlled by pre-
56
cipitation and evaporation. From east to west in North America, available

moisture decreases, influencing biome distribution. Humid regions along
the East Coast support forest biomes, including temperate coniferous for-
ests and temperate deciduous forest. West of the eastern forests is a biome
type of grasslands, including tall-grass prairie and short-grass steppe. In
this zone, there is less precipitation than evaporation. The ratio of precipi-
tation to evaporation is about 0.6 to 0.8 in the land that supports a tall-grass
prairie and 0.2 to 0.4 farther west, where a short-grass steppe is supported.
Beyond the short-grass steppe are shrubland and the deserts of the West.
Western Northern America is a mountainous country in which vegetation
zones reflect climatic changes on an altitudinal gradient. The vegetation in
the lowlands is characteristic of the regions (short-grass steppe in the east
side of Rocky Mountains, sagebrush cold semideserts in the Great Basin
between the Rocky Mountains and the Sierra Nevada, and grasslands in
California’s Central Valley west of the Sierra Nevada). Above the base re-
gions, the vegetation changes from shrub, woodland, or deciduous forest
to montane coniferous forests or alpine tundra. In Central America, from
Mexico to Panama where precipitation becomes ample and temperatures
are high, tropical rain forests and tropical seasonal forests occur.
Similar distributions of biomes along latitude and altitude can be found
in South America, Africa, and Eurasia. In general, the climate-induced pat-
terns of vegetation are influenced by latitude; the location of regions
within a continent, which affects the amount of moisture they receive; and
altitude, in which mountains modify the climate patterns. In addition,
other factors, such as fire and human disturbance, may influence distribu-
tions of biomes. For example, most grasslands require periodic fires for
maintenance, renewal, and elimination of incoming woody growth. Grass-
lands at one time covered about 42 percent of the land surface of the world.
Humans have converted much of that area into croplands.
Quantitative Relationships
Descriptive relationships can provide pictures of world vegetation distri-
butions along latitudinal and altitudinal gradients of temperature and
moisture. Ecologists in the past several decades have also sought quantita-
tive relationships between distributions of biomes and environmental fac-
tors. For example, when R. H. Whittaker plotted various types of biomes
on gradients of mean annual temperature and mean annual precipitation
in 1975, a global pattern emerged relating biomes to climatic variables. It
was shown that tropical rain-forest biomes are distributed in regions with
annual mean precipitation of 2,500 to 4,500 millimeters and annual mean
temperatures of 20 to 30 degrees Celsius. Tropical seasonal forest and sa-
57
vannas also occur in warm regions with precipitation of 1,500-2,500 milli-

meters and 500-1,500 millimeters per year, respectively. Temperate forests
occupy regions with annual temperature of 5 to 20 degrees Celsius and
precipitation exceeding 1,000 millimeters per year. This thermal zone can
support temperate rain forest when annual precipitation is more than 2,500
millimeters and temperate grassland when annual precipitation is below
750 millimeters. Temperate woodland occurs between temperate forests
and grasslands. Tundra and taiga are distributed in regions with annual
mean temperature below 3 degrees Celsius, whereas deserts occupy areas
with annual precipitation below 250 millimeters.
These relationships between climatic variables and biomes provide a
reasonable approximation of global vegetation patterns. Many types of
biomes intergrade with one another. Soil, exposure to fire, and regional cli-
mate can influence distributions of biomes in a given area.
Yiqi Luo
See also: Biomes: types; Chaparral; Deserts; Forests; Grasslands and prai-
ries; Habitats and biomes; Lakes and limnology; Marine biomes; Mediter-
ranean scrub; Mountain ecosystems; Old-growth forests; Rain forests; Rain
forests and the atmosphere; Rangeland; Reefs; Savannas and deciduous
tropical forests; Taiga; Tundra and high-altitude biomes; Wetlands.

Archibold, O. W. Ecology of World Vegetation. London: Chapman & Hall,
1995.
Smith, R. L., and T. M. Smith. Ecology and Field Biology. 6th ed. San Fran-
cisco: Benjamin Cummings, 2001.
Whittaker, R. H. Communities and Ecosystems. 2d ed. New York: Macmillan,
1975.
58
BIOMES: TYPES
Types of ecology: Biomes; Ecosystem ecology; Global ecology;
Theoretical ecology
Biomes are classified by geography, climate, temperature, precipitation, soil types,

and typical aggregates of flora and fauna. Although classification systems vary,
most agree that there are half a dozen major world biomes, which can be further
classified into minor biomes.
T emperature, precipitation, soil, and length of day affect the survival

and distribution of biome species. Species diversity within a biome
may increase its stability and capability to deliver natural services, includ-
ing enhancing the quality of the atmosphere, forming and protecting the
soil, controlling pests, and providing clean water, fuel, food, and drugs.
Major biomes include the temperate, tropical, and boreal forests; tundra;
deserts; grasslands; chaparral; and oceans.
Temperate Forests
The temperate forest biome occupies the so-called temperate zones in the
midlatitudes (from about 30 to 60 degrees north and south of the equator).
Temperate forests are found mainly in Europe, eastern North America, and
eastern China, and in narrow zones on the coasts of Australia, New Zea-
land, Tasmania, and the Pacific coasts of North and South America. Their
climates are characterized by high rainfall and temperatures that vary
from cold to mild.
Temperate forests contain primarily deciduous trees—including maple,
oak, hickory, and beechwood—and, secondarily, evergreen trees—includ-
ing pine, spruce, fir, and hemlock. Evergreen forests in some parts of the
Southern Hemisphere contain eucalyptus trees. The root systems of forest
trees help keep the soil rich. The soil quality and color are due to the action
of earthworms. Where these forests are frequently logged, soil runoff pol-
lutes streams, which reduces spawning habitat for fish. Raccoons, opos-
sums, bats, and squirrels are found in the trees. Deer and black bears roam
forest floors. During winter, small animals such as marmots and squirrels
burrow in the ground.
Tropical Forests
Tropical forests exist in frost-free areas between the Tropic of Cancer and
the Tropic of Capricorn. Temperatures range from warm to hot year-round.
59
Biomes: types
These forests are found in northern Australia, the East Indies, Southeast
Asia, equatorial Africa, and parts of Central America and northern South
America.
Tropical forests have high biological diversity and contain about 15 per-
cent of the world’s plant species. Animal life lives at each layer of tropical
forests. Nuts and fruits on the trees provide food for birds, monkeys, squir-
rels, and bats. Monkeys and sloths feed on tree leaves. Roots, seeds, leaves,
and fruit on the forest floor feed larger animals. Tropical forest trees pro-
duce rubber and hardwood, such as mahogany and rosewood. Deforesta-
tion for agriculture and pastures has caused reduction in plant and animal
diversity in these forests.
Boreal Forests
The boreal forest is a circumpolar Northern Hemisphere biome spread
across Russia, Scandinavia, Canada, and Alaska. The region is very cold.
Evergreen trees such as white spruce and black spruce dominate this zone,
which also contains larch, balsam, pine, fir, and some deciduous hard-
woods such as birch and aspen. The acidic needles from the evergreens
make the leaf litter that is changed into soil humus. The acidic soil limits
the plants that develop.
Biomes of the World
Tr o p i c o f
Cancer
Tr o p i c o f
Capricorn
Desert Tropical Rain Forest Temperate Grassland Taiga
Monsoon Savanna Temperate Forest Tundra
Mediterranean Mountain Polar
60
Biomes: types
Animals in boreal forests in- Terrestrial Biomes

clude deer, bears, and wolves. (percentages)
Birds in this zone include red-
tailed hawks, sapsuckers, grouse, Deciduous forest
7%
and nuthatches. Relatively few
animals emigrate from this hab- Other
itat during winter. Conifer seeds 15%
are the basic winter food. Rain Forest
12%
Tundra Coniferous Desert

About 5 percent of the earth’s forest 29%
12%
surface is covered with Arctic
tundra, and 3 percent with al- Grassland
pine tundra. The Arctic tundra 12% Tundra
13%
is the area of Europe, Asia, and
North America north of the
boreal coniferous forest zone,
where the soils remain frozen
most of the year. Arctic tundra
has a permanent frozen subsoil, called permafrost. Deep snow and low
temperatures slow the soil-forming process. The area is bounded by a 50
degrees Fahrenheit (122 degrees Celsius) circumpolar isotherm, known as
the summer isotherm. The cold temperature north of this line prevents
normal tree growth.
The tundra landscape is covered by mosses, lichens, and low shrubs,
which are eaten by caribou, reindeer, and musk oxen. Wolves eat these her-
bivores. Bears, foxes, and lemmings also live there. The most common Arctic
bird is the old squaw duck. Ptarmigans and eider ducks are also very com-
mon. Geese, falcons, and loons are some of the nesting birds of the area.
The alpine tundra, which exists at high altitude in all latitudes, is acted
upon by winds, cold temperatures, and snow. The plant growth is mostly
cushion- and mat-forming plants.
Deserts
The desert biome covers about one-seventh of the earth’s surface. Deserts
typically receive no more than 10 inches (25 centimeters) of rainfall per
year, and evaporation generally exceeds rainfall. Deserts are found around
the Tropic of Cancer and the Tropic of Capricorn. As warm air rises over
the equator, it cools and loses its water content. The dry air descends in the
two subtropical zones on each side of the equator; as it warms, it picks up
moisture, resulting in drying the land.
61
Biomes: types
Rainfall is a key agent in shaping the desert. The lack of sufficient plant
cover contributes to soil erosion during wind- and rainstorms. Some desert
plants—for example, the mesquite tree, which has roots that grow 40 feet
(13 meters) deep—obtain water from deep below the earth’s surface. Other
plants, such as the barrel cactus, store large amounts of water in their
leaves, roots, or stems. Some plants slow the loss of water by having tiny
leaves or shedding their leaves. Desert plants have very short growth peri-
ods, because they cannot grow during the long drought periods.
Grasslands
Grasslands cover about one-quarter of the earth’s surface and can be found
between forests and deserts. Treeless grasslands exist in parts of central
North America, Central America, and eastern South America that have be-
tween 10 and 40 inches (250-1,000 millimeters) of erratic rainfall per year.
The climate has a high rate of evaporation and periodic major droughts.
Grasslands are subject to fire.
Some grassland plants survive droughts by growing deep roots, while
others survive by being dormant. Grass seeds feed the lizards and rodents
that become the food for hawks and eagles. Large animals in this biome in-
clude bison, coyotes, mule deer, and wolves. The grasslands produce more
food than any other biome. Overgrazing, inefficient agricultural practices,
and mining destroy the natural stability and fertility of these lands, result-
ing in reduced carrying capacity, water pollution, and soil erosion. Diverse
natural grasslands appear to be more capable of surviving drought than
are simplified manipulated grass systems. This may be due to slower soil
mineralization and nitrogen turnover of plant residues in the simplified
system.
Savannas are open grasslands containing deciduous trees and shrubs.
They are near the equator and are associated with deserts. Grasses there
grow in clumps and do not form a continuous layer.
Chaparral
The chaparral, or Mediterranean, biome is found in the Mediterranean Ba-
sin, California, parts of Australia, middle Chile, and the Cape Province of
South America. This region has a climate of wet winters and summer
drought. The plants have tough, leathery leaves and may have thorns. Re-
gional fires clear the area of dense and dead vegetation. The seeds from
some plants, such as the California manzanita and South African fire lily,
are protected by the soil during a fire and later germinate and rapidly grow
to form new plants. Vegetation dwarfing occurs as a result of the severe
summer drought and extreme climate changes.
62
Biomes: types
Oceans
The ocean biome covers more than 70 percent of the earth’s surface and in-
cludes 90 percent of its volume. Oceans have four zones. The intertidal
zone is shallow and lies at the land’s edge. The continental shelf, which be-
gins where the intertidal zone ends, is a plain that slopes gently seaward.
The neritic zone (continental slope) begins at a depth of about 600 feet (180
meters), where the gradual slant of the continental shelf becomes a sharp
tilt toward the ocean floor, plunging about 12,000 feet (3,660 meters) to the
ocean bottom. This abyssal zone is so deep that it does not have light.
Plankton are animals that float in the ocean. They include algae and
copepods, which are microscopic crustaceans. Jellyfish and animal larva
are also considered plankton. The nekton are animals that move freely
through the water by means of their muscles. These include fish, whales,
and squid. The benthos are animals that are attached to or crawl along the
ocean’s floor. Clams are examples of benthos. Bacteria decompose the dead
organic materials on the ocean floor.
The circulation of materials from the ocean’s floor to the surface is
caused by winds and water temperature. Runoff from the land contains
pollutants such as pesticides, nitrogen fertilizers, and animal wastes.
Rivers carry loose soil to the ocean, where it builds up the bottom areas.
Overfishing has caused fisheries to collapse in every world sector.
Human Impact on Biomes

Human interaction with biomes has increased biological invasions, re-
duced species biodiversity, changed the quality of land and water re-
sources, and caused the proliferation of toxic compounds. Managed care of
biomes may not be capable of undoing these problems.
Ronald J. Raven
See also: Biomes: determinants; Chaparral; Deserts; Forests; Grasslands

and prairies; Habitats and biomes; Lakes and limnology; Marine biomes;
Mediterranean scrub; Mountain ecosystems; Old-growth forests; Rain for-
ests; Rain forests and the atmosphere; Rangeland; Reefs; Savannas and de-
ciduous tropical forests; Taiga; Tundra and high-altitude biomes; Wet-
lands.

Food and Agriculture Organization of the United Nations. State of the
World’s Forests, 2001. Rome: Author, 2001.
Gawthorp, Daniel, and David Suzuki. Vanishing Halo: Saving the Boreal For-
est. Seattle: Mountaineers, 1999.
63
Biomes: types
Linsenmair, K. E., ed. Tropical Forest Canopies: Ecology and Management. Lon-
don: Kluwer Academic, 2001.
Prager, Ellen J., with Cynthia A. Earle. The Oceans. New York: McGraw-
Hill, 2000.
Solbrig, Otto Thomas, E. Medina, and J. F. Silva, eds. Biodiversity and Sa-
vanna Ecosystem Processes: A Global Perspective. New York: Springer,
1996.
64
BIOPESTICIDES
Types of ecology: Agricultural ecology; Ecotoxicology
Biopesticides are biological agents, such as viruses, bacteria, fungi, mites, and
other organisms used to control insect and weed pests in an environmentally and
ecologically friendly manner.
B iopesticides allow biologically based, rather than chemically based,

control of pests. Pests are any unwanted animal, plant, or microorgan-
ism. When the environment provides no natural resistance to a pest and
when no natural antagonists are present, pests can run rampant. For exam-
ple, spread of the fungus Endothia parasitica, which entered New York in
1904, caused the nearly complete destruction of the American chestnut tree
because no natural control was present. Viruses, bacteria, fungi, protozoa,
mites, insects, and flowers have all been used as biopesticides.
Advantages and Disadvantages

Many plants and animals are protected from pests by passive means. For
example, plant rotation is a traditional method of insect and disease pro-
tection that is achieved by removing the host plant long enough to reduce a
region’s pathogen and pest populations. Biopesticides have several signifi-
cant advantages over commercial pesticides. They appear to be ecologi-
cally safer than commercial pesticides because they do not accumulate in
the food chain. Some biopesticides provide persistent control, as more than
a single mutation is required to adapt to them and because they can be-
come an integral part of a pest’s life cycle. In addition, biopesticides have
slight effects on ecological balances because they do not affect nontarget
species. Finally, biopesticides are compatible with other control agents.
The major drawbacks to using biopesticides are the time required for
them to kill their targets and the inefficiency with which they work. Also, if
the organism being used as a biopesticide is a nonnative species, it may
cause unforeseen damage to the local ecosystem.
Viruses and Bacteria

Viruses have been developed against insect pests such as Lepidoptera (but-
terflies and moths), Hymenoptera (bees, wasps, and ants), and Dipterans
(flies). Gypsy moths and tent caterpillars, for example, periodically suffer
from epidemic virus infestations, which could be exploited and encour-
aged.
65
Biopesticides
Many commensal microorganisms (microorganisms that live on or in

other organisms causing no direct benefit or harm) that occur on plant
roots and leaves can passively protect plants against microbial pests by
competitive exclusion (that is, simply crowding them out). Bacillus cereus
has been used as an inoculum on soybean seeds to prevent infection by
fungal pathogens in the genus Cercospora. Some microorganisms used as
biopesticides produce antibiotics, but the major mechanism in most cases
seems to be competitive exclusion. For example, Agrobacterium radiobacter
antagonizes Agrobacterium tumefaciens, which causes the disease crown
gall. Species of two bacterial genera–Bacillus and Streptomyces—when
added as biopesticides to soil, help control the damping-off disease of cu-
cumbers, peas, and lettuce caused by Rhizoctonia solani. Bacillus subtilis
added to plant tissue also controls stem rot and wilt rot caused by species
of the fungus Fusarium. Mycobacteria species produce cellulose-degrading
enzymes, and their addition to young seedlings helps control fungal infec-
tion by species of Pythium, Rhizoctonia, and Fusarium. Species of Bacillus
and Pseudomonas produce enzymes that dissolve fungal cell walls.
Bacillus thuringiensis Toxins

The best examples of microbial insecticides are Bacillus thuringiensis (B.t.)
toxins, which were first used in 1901. They have had widespread commer-
cial production and use since the 1960’s and have been successfully tested
on 140 insects, including mosquitoes. Insecticidal endotoxins are produced
by B.t. during sporulation, and exotoxins are contained in crystalline
parasporal protein bodies. These protein crystals are insoluble in water but
readily dissolve in an insect’s gut. Once dissolved, the proteolytic enzymes
paralyze the gut. Spores that have been consumed germinate and kill the
insect. Bacillus popilliae is a related bacterium that produces an insecticidal
spore that has been used to control Japanese beetles, a corn pest.
The gene for the B.t. toxin has also been inserted into the genomes
of cotton and corn, producing genetically modified, or GM, plants that
produce their own B.t. toxin. GM cotton and B.t. corn both express the gene
in their roots, which provides them with protection from root worms. Ecol-
ogists and environmentalists have expressed concern that constantly ex-
posing pests to B.t. will cause insects to develop resistance to the toxin. In
such a scenario, the effectiveness of traditionally applied B.t. would de-
crease.
Fungi and Protozoa

Saprophytic fungi can compete with pathogenic fungi. There are several
examples of fungi used as biopesticides, such as Gliocladium virens, Tri-
66
Biopesticides
choderma hamatum, Trichoderma harzianum, Trichoderma viride, and Talaromyces

flavus. For example, Trichoderma species compete with pathogenic species
of Verticillium and Fusarium. Peniophora gigantea antagonizes the pine patho-
gen Heterobasidion annosum by three mechanisms: It prevents the pathogen
from colonizing stumps and traveling down into the root zone, it prevents
the pathogen from traveling between infected and uninfected trees along
interconnected roots, and it prevents the pathogen from growing up to
stump surfaces and sporulating.
Nematodes are pests that interfere with commercial button mushroom
(Agaricus bisporus) production. Several types of nematode-trapping fungi
can be used as biopesticides to trap, kill, and digest the nematode pests.
The fungi produce constricting and nonconstricting rings, sticky append-
ages, and spores, which attach to the nematodes. The most common of the
nematode-trapping fungi are Arthrobotrys oligospora, Arthrobotrys conoides,
Dactylaria candida, and Meria coniospora.
Protozoa have occasionally been used as biopesticide agents, but their
use has suffered because of slow growth and the complex culture condi-
tions associated with their commercial production.
Mites, Insects, and Flowers

Well-known “terminator” bugs include praying mantis and ladybugs as
well as decollate snails, which eat the common brown garden snail. Fleas,
grubs, beetles, and grasshoppers often have natural nematode species that
prey on them, which can be used as biocontrol agents. Predaceous mites
are used as a biopesticide to protect cotton from other insect pests such as
the boll weevil. Parasitic wasps of the genus Encarsia, especially E. formosa,
prey on whiteflies, as does Delphastus pusillus, a small, black ladybird
beetle.
Dalmatian and Persian insect powders contain pyrethrins, which are
toxic insecticidal compounds produced in chrysanthemum flowers. Syn-
thetic versions of these naturally occurring compounds are found in prod-
ucts used to control head lice.
Mark S. Coyne, updated by Elizabeth Slocum
See also: Biomagnification; Food chains and webs; Genetically modified

foods; Integrated pest management; Multiple-use approach; Pesticides;
Pollution effects; Soil; Soil contamination.

Carozzi, Nadine, and Michael Koziel, eds. Advances in Insect Control: The
Role of Transgenic Plants. Bristol, Pa.: Taylor & Francis, 1997.
67
Biopesticides
Deacon, J. W. Microbial Control of Plant Pests and Diseases. Washington, D.C.:

American Society for Microbiology, 1983.
Hall, Franklin R., and Julius J. Menn, eds. Biopesticides: Use and Delivery.
Totowa, N.J.: Humana Press, 1999.
Hokkanen, Heikki M. T., and James M. Lynch, eds. Biological Control: Bene-
fits and Risks. New York: Cambridge University Press, 1995.
68
BIOSPHERE CONCEPT
Types of ecology: Global ecology; Theoretical ecology
The term “biosphere” was coined in the nineteenth century by Austrian geologist
Eduard Suess in reference to the 20-kilometer-thick zone extending from the floor
of the oceans to the top of mountains, within which all life on earth exists. Thought
to be more than 3.5 billion years old, the biosphere supports nearly one dozen
biomes, regions of climatic conditions within which distinct biotic communities
reside.
C ompounds of hydrogen, oxygen, carbon, nitrogen, potassium, and

sulfur are cycled among the four major spheres, one of which is the
biosphere, to make the materials that are essential to the existence of life.
The other spheres are the lithosphere, the outer part of the earth; the atmo-
sphere, the whole mass of air surrounding the earth; and the hydrosphere,
the aqueous vapor of the atmosphere, sometimes defined as including the
earth’s bodies of water.
The Water Cycle

The most critical of these compounds is water, and its movement among
the spheres is called the hydrologic cycle. Dissolved water in the at-
mosphere condenses to form clouds, rain, and snow. The annual pre-
cipitation for any region is one of the major factors in determining the
terrestrial biome that can exist. The precipitation takes various paths
leading to the formation of lakes and rivers. These flowing waters in-
teract with the lithosphere (the outer part of the earth’s crust) to dis-
solve chemicals as they flow to the oceans. Evaporation of water from
the oceans then supplies most of the moisture in the atmosphere. This cy-
cle continually moves water among the various terrestrial and oceanic
biomes.
Solar Energy
The biosphere is also dependent upon the energy that is transferred from
the various spheres. Solar energy is the basis for almost all life. Light enters
the biosphere as the essential energy source for photosynthesis. Plants take
in carbon dioxide, water, and light energy, which is converted via photo-
synthesis into chemical energy in the form of sugars and other organic
molecules. Oxygen is generated as a by-product. Most animal life reverses
69
Biosphere concept
This composite image of the earth’s biosphere shows the planet’s heaviest vegeta-
tive biomass in the dark sections, known to be rain forests. The combination of in-
tense sunlight for twelve hours per day, warm temperatures throughout the year,
and considerable rainfall make tropical rain forests the most productive ecosystems
on land. (NASA)
this process during respiration, as chemical energy is released to do work

by the oxidation of organic molecules to produce carbon dioxide and
water.
Incoming solar energy also interacts dramatically with the water cycle
and the worldwide distribution of biomes. Because of the earth’s curva-
ture, the equatorial regions receive a greater amount of solar heat than the
polar regions. Convective movements in the atmosphere—such as winds,
high- and low-pressure systems, and weather fronts—and the hydro-
sphere—such as water currents—are generated during the redistribution
of this heat. The weather patterns and climates of earth are a response to
these energy shifts. Earth’s various climates are defined by the mean an-
nual temperature and the mean annual precipitation.
Toby R. Stewart and Dion Stewart
See also: Biodiversity; Biomes: determinants; Biomes: types; Ecology: defi-

nition; Ecosystems: definition and history; Ecosystems: studies; Geochem-
ical cycles; Global warming; Greenhouse effect; Habitats and biomes;
Hydrologic cycle; Ozone depletion and ozone holes; Rain forests and the
atmosphere.
70
Biosphere concept

McNeely, Jeffrey A. Conserving the World’s Biological Diversity. Washington,
D.C.: International Union for Conservation of Nature and Natural Re-
sources, 1990.
Smith, Vaclav. Cycles of Life: Civilization and the Biosphere. New York: W. H.
Freeman, 2000.
Vernadskii, V. I. The Biosphere. Translated by Mark A. S. McMenamin. New
York: Copernicus, 1998.
Weiner, Jonathon. The Next One Hundred Years: Shaping the Fate of Our Living
Earth. New York: Bantam Books, 1991.
Wilson, Edward O., ed. Biodiversity. Washington, D.C.: National Academy
Press, 1988.
71
CAMOUFLAGE
Type of ecology: Physiological ecology
Both predatory and prey species use camouflage to minimize the chance that their
presence will be detected. Although camouflage is often thought of as being exclu-
sively a visual phenomenon, for it to work, it must occur in all sensory modalities.
C rypsis is the art of remaining hidden. Camouflage is usually thought of

as color matching: a green aphid, for example, is likely to go unnoticed
while feeding on a green leaf. Background matching, or cryptic coloration,
is indeed the most common form of camouflage, but most crypsis involves
far more than matching a single color. Very small animals such as aphids
can get away with using a single camouflage color because they are much
smaller than the plants on which they spend their entire lives: They need to
match only one thing. Most animals, however—even most insects—are
significantly larger than aphids and are likely to spend time in more than
one place. Their camouflage must be more sophisticated if it is to be useful.
Disruptive Coloration
If a large organism is to remain undetected, it must be camouflaged with
respect to an entire scene. One way to do this is with the use of disruptive
coloration, that is, the use of stripes, spots, or patches of color for camou-
flage. Disruptive coloration can involve large color patches, as on a pinto
pony, a tabby cat, or a diamond-backed rattlesnake, or may involve tiny
variations of color on each scale, feather, or hair. Many brownish or grayish
mammals actually have what is called agouti coloring, with three different
colors appearing on each hair.
The irregular borders of multiple color patches on an animal’s body
help to obscure its outline against an irregular and multicolored back-
ground, just like the blotchy greens and browns on military uniforms. An
animal that has a mix of browns in its fur, feathers, skin or scales, for exam-
ple, will blend into a forest or even an open desert or tundra much better
than one that is a single solid color. Even the black-and-white stripes of ze-
bras, which seem so striking, act as a form of disruptive coloration: From
far away, and especially to an animal such as a lion, which does not have
good color vision, the stripes of zebras help them blend into the tall, wavy
grasses of the savannah.
Countershading is another form of crypsis, involving differently col-
ored patches. Countershaded animals appear dark when viewed from
72
Camouflage
above and light when viewed from underneath. Animals with counter-
shading include orca whales with their black backs and white bellies, pen-
guins, blue jays, bullfrogs, and weasels. Countershading works and is
found as camouflage in so many kinds of animals because no matter where
one lives—a desert, a forest, a meadow, or an ocean—the sun shines from
above. When looking up toward the sun and sky, dark things stand out and
light colors blend in; when looking down toward the ground or the ocean
floor, light colors stand out and dark colors blend in. Predators that are
countershaded can thus approach their prey with equal stealth from either
above or below; likewise, prey species that are countershaded will be
equally hard to find whether a predator is searching from on high or from
underneath. Countershading and other forms of disruptive coloration can
occur in the same organism, so that dark spots, blotches, or stripes appear
on top while paler ones appear below.
Mimicry
Another way of remaining undetected in a complex scene is by using pro-
tective mimicry, the ability to mimic an inanimate object in both color and
form. Some insects look like thorns on plant stems; others look like leaves
or twigs or flowers. Some insects, frogs, and fish look like rocks, lichens, or
corals. Sea lions, sea dragons, and even eels can look like floating kelp or
other forms of seaweed.
Zebras’ stripes not only help them blend into their grassland habitats but also
make it difficult for predators to pick out a single individual for attack. (Digital
Stock)
73
Camouflage
Some animals may not look much like the objects around them but will
disguise themselves by attaching pieces of plants or sand or other debris to
their body. Some caterpillars use silk to tie bits of flowers and leaves to
their bodies; others use saliva as a glue. Some crabs glue broken bits of
shell and coral to their exoskeletons. By using local materials to camou-
flage itself, an animal can ensure that it matches the background and can
even change its disguise as it moves from one area into another.
Transparency
Being transparent is another way to match whatever background happens
to be present. Many marine invertebrates such as worms, jellyfish, and
shrimp, are completely transparent. Complete transparency is less com-
mon among land animals, but some land invertebrates have transparent
body parts, such as their wings, allowing them to break up the outline of
their body and blend into whatever happens to be in the immediate back-
ground.
Predation and Prey

Behavior is an important factor in the success or lack of success of any form
of crypsis. For example, not even disruptive camouflage can hide some-
thing that is moving quickly with respect to its background. Predatory spe-
cies that rely on speed or stamina to outrun, outswim, or outfly their prey
therefore have little use for camouflage. On the other hand, so-called sit-
and-wait predators (such as boa constrictors or praying mantises) must be
virtually perfectly camouflaged in order to remain undetected while their
prey approach to within grabbing distance. In between are the stealth
hunters, which sneak up on their prey before making a final high-speed at-
tack; such animals must be camouflaged and slow-moving when out of at-
tack range but do not have to be camouflaged or slow when at close range.
Like predators, prey species that rely on rapid escape maneuvers do not
often exhibit camouflage coloration, while prey species that cannot rely on
efficient escape tactics must, instead, rely on camouflage to avoid being
seen in the first place. Prey species that can move quickly but not as quickly
as their predators must detect their predators before their predators detect
them, and then they must remain absolutely still until the danger is passed.
Life Stages
Some species use different strategies as they go through different stages in
life. In many altricial species (species with dependent young that require
extended parental care of the offspring), the eggs and/or young are cam-
ouflaged, even though the adults are not; the temporary spots on deer
74
Camouflage
fawns and mountain lion cubs are examples. In other species, nesting or
brooding females may be camouflaged while the adult males retain their
gaudy plumage or attention-getting behaviors; the changing seasonal pat-
terns of color and behavior of ducks and songbirds provide examples here.
Some species may be toxic and gaudy during one stage of life, yet tasty and
cryptic during another.
Multisensory Camouflage
Finally, although camouflage is usually thought of as a visual phenome-
non, crypsis is important in every sensory modality. If a prey animal is vir-
tually invisible to its predators but puts out a sound, a scent, or a vibration
that makes it easy to locate, visual crypsis alone is useless. For successful
protection, prey species must be cryptic in whatever sensory modalities
their predators use for hunting. Likewise, for successful hunting, preda-
tory species must be cryptic in whatever sensory modalities their prey use
to detect danger. For most species of both predator and prey, this means be-
ing camouflaged or blending into the background in several sensory mo-
dalities all at once.
Linda Mealey
See also: Adaptations and their mechanisms; Bioluminescence; Defense

mechanisms; Displays; Metabolites; Mimicry; Pollination.

Dettner, K., and C. Liepert “Chemical Mimicry and Camouflage.” Annual
Review of Entomology 39 (1994): 129-154.
Ortolani, Alessia. “Spots, Stripes, Tail Tips, and Dark Eyes: Predicting the
Function of Carnivore Colour Patterns Using the Comparative
Method.” Biological Journal of the Linnean Society 67, no. 4 (August, 1999):
433-476.
Owen, Denis. Camouflage and Mimicry. Chicago: University of Chicago
Press, 1980.
Ramachandran, V. S., et.al. “Rapid Adaptive Camouflage in Tropical
Flounders.” Nature 379, no. 6568 (1996): 815-818.
Wicksten, Mary K. “Decorator Crabs.” Scientific American 242, no. 2 (Febru-
ary, 1980): 146-154.
75
CHAPARRAL
Types of ecology: Biomes; Ecosystem ecology
Chaparral is the name of a major ecosystem (or biome) found in areas with moist,
cool to cold winters and long, dry hot summers (Mediterranean climate).
C haparral ecosystems with different names occur in the Mediterra-

nean, South Africa, Chile, Australia, and Mexico. The word “chapar-
ral” is a colloquial adaptation of the original Mexican name, chaparro.
Chaparral communities in other parts of the world have the same basic
characteristics and very similar adaptations; this article focuses on the
chaparral of California and the American Southwest.
Chaparral is an interesting and unique ecosystem. It is an elfin (stunted)
forest dependent on a fire ecology, and its adaptations to a harsh and vari-
able climate are remarkable. The chaparral’s geology, latitude, altitude,
and climate are all related and have played a role in its formation.
Image Not Available
76
Chaparral
Location
In California, the chaparral is located mainly along the central and south-
ern coastal areas, primarily between elevations of 500 and 2,500 feet. It is
also found in some areas of the Sierra Nevada foothills, one hundred miles
or more inland, and in the lower elevations of some other interior moun-
tains. The geology of most of the areas where this ecosystem occurs is be-
lieved to have started with massive upheavals from half a million to ten
thousand years ago. The most common substrate was granite. Relentless
disintegration resulted in rocky and sandy debris which would allow in-
creasing amounts of plant life to grow. As the organic matter became more
abundant, its debris (leaves, twigs, and decaying dead plants) became
more and more mixed into the materials of the granite decomposition. This
resulted in a rich, sandy loam.
Chaparral Flora
Because of the cool, moist winters and dry, hot summers, plants evolved to
survive these marked changes. In most of the chaparral, there are quite ex-
treme diurnal temperature changes, with fluctuations of fifty to sixty de-
grees or more. Compounding these harsh conditions are frequent strong,
dry winds, often reaching forty miles per hour.
The plants that have become the residents of the chaparral are mainly
shrubby, small-leaved evergreens with leathery, thick stems. Shrubs pre-
dominate, but there are also small trees and in many areas abundant wild-
flowers. All plants are adapted to conserve water. There is little humus
in the soil, which is relatively nutrient-poor. The sandy nature of the
soil and the variable periods of dry and sudden rain can cause a quick
runoff.
The predominant plants are between three and nine feet in height, with
some trees being taller. They hug close to the ground to provide shade. The
ratio of the surface area of the leaves and stems to their body mass is re-
duced, and they tend to have thick, heat-resistant surfaces. Some of the
plants are capable of turning their leaves so the edges face the sun, which
cuts down the warming effect on their surfaces. All these mechanisms
greatly reduce evaporative water loss. Most of the bushes and trees also
have unusually long tap roots. A three-foot plant might have a tap root that
goes ten or more feet below the surface, enabling it to get more water and
nutrients.
The most common plant in the chaparral is the greasewood-chamise
bush (Adenostoma fasciculatum). Others that predominate are the christmas-
berry-toyon bush (Photinia arbutifolia), the scrub oak (Quercus dumosa), the
yucca (Yucca whipplei), and the hoary manzanita (Arctostaphylos canescens).
77
Chaparral
The chamise is characterized by numerous small, club-shaped leaves with

a waxy substance that protects them from drying. When there is a fire, the
chamise burns with an intense heat and creates a very black smoke (hence
the name “greasewood”).
The Role of Fire

The role of fire in the maintenance and regeneration of the chaparral is of
paramount importance. The hot, dry summer weather, often fanned by
winds, makes the chaparral very prone to fires. Because of the high fuel
content in the dense plants, with their waxy and oily components, these
fires are very intense and can spread rapidly. Fire is necessary to clear ex-
cess growth and allow new seeds to germinate. Indeed, several of the key
species need fire to release their seeds or they will not germinate. The
amount and distribution of the canopy fuel can have a marked effect on
regrowth and even spatial variation. Naturally recurring fires are usually
good for germination.
However, unusually intense fires, often from years of fire suppression,
may do harm by damaging the plants severely. In much of the chaparral,
human interference has caused this suppression, creating a difficult para-
dox, since many people now reside in the chaparral and chaparral fires can
spread very rapidly, especially with strong winds. Conversely, in some ar-
eas where fire has been controlled, the chaparral has been retreating. A
good example of this retreat has occurred on the southern slopes of Mount
Tamalpais, a mountain just north of San Francisco.
Chaparral Fauna
A wide variety of reptiles, birds, and mammals make the chaparral their
home. They have developed adaptations to survive and thrive in this harsh
environment. Several species of skinks, lizards, and a variety of snakes, in-
cluding gopher snakes, the California king snake, and both the red dia-
mond and western rattlesnakes are residents. There are dozens of birds,
from several species of hummingbirds to the large birds: the turkey vul-
ture, barn owl, roadrunner, and golden eagle. There are many species of ro-
dents, including kangaroo rats, chipmunks and gophers. The variety of
medium to large mammals of common interest is impressive and includes
the coyote, gray fox, badger, lynx, bobcat, mountain lion, and mule deer.
C. Mervyn Rasmussen
See also: Biomes: determinants; Biomes: types; Deserts; Forests; Grass-

lands and prairies; Habitats and biomes; Lakes and limnology; Marine
biomes; Mediterranean scrub; Mountain ecosystems; Old-growth forests;
78
Chaparral
Rain forests; Rain forests and the atmosphere; Rangeland; Reefs; Savannas
and deciduous tropical forests; Taiga; Tundra and high-altitude biomes;
Wetlands.

Brown, O. E., and R. A. Minnich. “Fire and Changes in the Creosote Bush
Scrub of the Western Sonoran Desert, California.” American Midland
Naturalist 116, no. 2 (1986): 411-422.
Collis, P. H., ed. Dictionary of Ecology and the Environment. 3d ed. Chicago:
Fitzroy Dearborn, 1998.
Head, W. S. The California Chaparral: An Elfin Forest. 1972. Reprint. Happy
Camp, Calif.: Naturegraph, 1998.
Jasson-Holt, Sophie. Unfold the Chaparral. San Francisco: San Francisco
State University Chapbook, 1996.
Odion, Dennis C., and Frank W. Davis. “Fire, Soil Heating, and the Forma-
tion of Vegetation Patterns in Chaparral.” Ecological Monographs 70,
no. 1 (2000): 149-169.
79
CLINES, HYBRID ZONES, AND
INTROGRESSION
Types of ecology: Population ecology; Speciation
A cline is a genetic variation in the characteristics of populations of the same spe-

cies that results from a variation in the geographical area that it occupies. Hybrid
zones are areas with populations of a species composed of individuals with charac-
teristics of one or more species that have interbred. Introgression is speciation that
occurs when the genes of one species are incorporated into the gene pool of another
as the result of successful hybridization.
G ene flow among populations tends to increase the similarity of char-

acters among all the demes (local populations) of a species. Natural
selection has the opposite effect: It tends to make every deme uniquely spe-
cialized for its specific habitat. Clines are one possible result of these two
opposing forces; a cline is a phenomenon in which a genetic variation oc-
curs that is caused by a difference in geographical habitat. Each species is
continuously adjusting its gene pool to ensure that the species survives in
the face of an environment that is continuously changing.
Comparing the characteristics of the demes of a single species usually
will reveal that they are not identical. The greater the distance between the
demes, the greater the differences between them will be. The grass frogs in
Wisconsin differ from the grass frogs in Texas more than they differ from
those in Michigan. On the average, the song sparrows of Alaska are
heavier and have darker coloration than those in California. These phe-
nomena, in which a single character shows a gradient of change across a
geographical area, are called clines.
North-South Clines
Many birds and mammals exhibit north-south clines in average body size
and weight, being larger and heavier in the colder climate farther north
and smaller and lighter in warmer climates to the south. In the same way,
many mammalian species show north-south clines in the sizes of body ex-
tremities such as tails and ears, these parts being smaller in northern
demes and larger in southern demes. Increase in average body size with in-
creasing cold is such a common observation that it has been codified as
Bergmann’s rule. The tendency toward shorter and smaller extremities in
colder climates and longer and larger ones in warmer climates is known as
80
Clines, hybrid zones, and introgression
Allen’s rule. The trend toward lighter colors in southern climates and
darker shades in northern climates has been designated Gloger’s rule. The
zebra, for example, shows a cline in the amount of striping on the legs. The
northernmost races are fully leg-striped, and the striping diminishes to-
ward the southern latitudes of Africa; this appears to be an example of
Gloger’s rule.
Another example of a cline, which does not fit any of the biogeo-
graphical rules mentioned, is the number of eggs laid per reproductive ef-
fort (the clutch size) by the European robin: This number is larger in north-
ern Europe than it is for the same species in northern Africa. Other birds,
such as the crossbill and raven, which have wide distribution in the
Holarctic realm, show a clutch-size cline that reveals a larger clutch size in
lower latitudes. The manifestation of such clines in clutch size is a conse-
quence of the interplay of two different reproductive strategies that may
give a species a competitive advantage in a given environment. The stabil-
ity of the environment is what elicits the appropriate strategy.
In unstable environments, such as those in the temperate zone, where
there may occur sudden variations in weather and extremes between sea-
sons, a species needs to reproduce rapidly and build its numbers quickly to
take advantage of the favorable warm seasons to ensure survival of the
species during the harsh, unfavorable conditions of winter. This strategy is
known as r strategy (r stands for the rate of increase). In the tropics, the cli-
mate is more equable throughout the year. The environment, however, can
only support a limited number of individuals throughout the year. This
number is called the carrying capacity. When carrying capacity is reached,
competition for resources increases, and the reproductive effort is reduced
to maintain the population at the carrying capacity. This is called K strat-
egy, with K standing for carrying capacity.
In birds, clutch size tends to be inversely proportional to the climatic
stability of the habitat: In temperate climates, more energy is directed to in-
crease the reproductive rate. In the tropics, the carrying capacity is more
important, resulting in a reduced reproductive rate. In the apparent con-
tradiction of the crossbills and ravens, it may be the harshness of the habi-
tat at higher latitudes that limits the resources available for successfully
fledging a larger number of young.
Frog Clines
The cline exhibited by the common grass frog is one of the best known of
all the examples of this phenomenon. It has the greatest range, occupies the
widest array of habitats, and possesses the greatest amount of morphologi-
cal variability of any frog species. This variability and adaptation are not
81
haphazard. The species includes a number of temperature-adapted demes,

varying from north to south. These adaptations involve the departmental
processes from egg to larva. The northernmost demes have larger eggs that
develop faster at lower temperatures than those of the southernmost
demes. These physiological differences are so marked that matings be-
tween individuals from the extreme ends of the cline result in abnormal
larvae or offspring that are inviable (cannot survive) even at a temperature
that is average for the cline region.
Leopard frogs from Vermont can interbreed readily with ones from
New Jersey. Those in New Jersey can hybridize readily with those in the
Carolinas, and those in turn with those in Georgia. Yet, hybrids of Vermont
demes and Florida demes are usually abnormal and inviable. Thus, it ap-
pears that the Vermont gene pool has been selected for a rate of develop-
ment that corresponds to a lower environmental temperature. The gene
pool of the Florida race has a rate of development that is slower at a higher
average temperature. The mixture of the genetic makeup of the northern
and southern races is so discordant that it fails to regulate characteristic
rates of development at any sublethal temperature, so the resulting em-
bryo dies before it becomes a tadpole.
There are two primary reasons why characters within a species may
show clinal variation. First, if gene flow occurs between nearby demes of a
population, the gene pools of demes that are close to one another will share
more alleles than the gene pools of populations that are far apart. Second,
environmental factors, such as annual climate, vary along gradients that
can be defined longitudinally, latitudinally, or altitudinally. Because these
environmental components act as selective pressures, the phenotypic char-
acters that are best adapted to such pressures will also vary in a gradient.
Hybridization, Hybrid Swards, Introgressive Demes

Hybridization is the process whereby individuals of different species pro-
duce offspring. A hybrid zone is an area occupied by interbreeding species.
Partial species can and do develop on the way to becoming new species as
products of hybridization. Natural hybridization and gene flow can take
place between biological species no matter how sterile most of the hybrid
offspring may be. As long as the mechanisms that prevent free exchange of
genes between populations can be penetrated, there is the potential for a
new species to develop. Because the parental species has a tendency to be
replaced by the hybrid types if natural selection favors them, hybridization
can be a threat to the integrity of the parental species as a distinct entity.
Hybridization between different species leads to various and unpre-
dictable results. Any time that hybridization occurs, the isolation mecha-
82
nisms of populations are overcome, forming bridging populations. Such

connecting demes of hybrid origin fall into one of two general categories:
hybrid swarms or introgressive demes. The formation of these types of
demes reverses the process of speciation and changes the formerly distinct
species into a complex mixture of highly variable individuals that are the
products of the segregation and independent assortment of traits. This is
the primary advantage of sexual reproduction: to produce variation in the
population that is acted upon by natural selection over time. It cannot be
overemphasized that hybrid swarms and introgressive demes are highly
variable.
The environmental conditions that contour animal communities have
endured for a very long time. In long-lived communities, every available
niche has been filled by well-adapted species. When populations with new
adaptive characteristics occur, there is no niche for them to occupy, so they
usually die out. In contrast, when such communities are disturbed, the
parity among their component species is upset, which gives new variants
an opportunity to become established.
Hybrid swarms can be observed in nature by the careful investigator.
The hybrid swarm forms in a disturbed habitat, where hybrid individuals
backcross with the parental types to form a third population, which result
from the migration of the genes of one population into the other. Such a
population is designated an introgressive population. The progeny of such
populations resemble the parent species, but the variations are in the direc-
tion of one parental species or the other. If introgression is extensive
enough, it may eradicate the morphological and ecological distinctions of
the parental types. The parental types become rarer and rarer, until they
are no longer the representatives of the species.
There appear to be three reasons that first-generation hybrids occurring
naturally are more likely to form offspring by backcrossing to one of the pa-
rental species than by mating with each other. Primarily, the hybrids are al-
ways rarer than the parents. Second, the parental individuals are so much
more fertile than the hybrids that many more parental gametes are avail-
able than hybrid ones. Finally, backcross progeny, since they contain pri-
marily parentally derived genes, are more likely to be well adapted to the
habitat in which they originated than are the purely hybrid individuals.
Introgression
Thus, the most likely result of hybridization is backcrossing to one of the
parental species. Genotypes containing the most parental genes usually
have the selective advantage, and the fact that they contain a few chromo-
somal segments from another species gives them unique characteristics
83
that may also be advantageous. This sequence of events—hybridization,

backcrossing, and stabilization of backcross types—is known as intro-
gression. Hybrid swarms are interesting phenomena, but they are unlikely
to be of evolutionary significance except through introgression.
There are many examples of introgression among plants, but examples
of introgression in animals are not common. Those that have been demon-
strated are usually associated with the domestication of livestock. In the
Himalayan region of Asia, there exists a relative of cattle, the yak, which is
also domesticated. Many of the herds of cattle found along the western
edge of the Himalayas, in central Asia, contain characteristics that clearly
are derived from the gene pool of the yak. Many of these characteristics are
manifested as adaptations to the harsh climatic conditions in this region.
In western Canada, there has been a modest introgression of the genes
of the American bison into the gene pool of strains of range cattle. The
bisonlike characters incorporated into beef cattle created a new breed
called the beefalo, which exhibits such characteristics as greater body mus-
culature, lower fat content of the flesh, and great efficiency in the utiliza-
tion of range forage. A beefalo steer is ready for market in only eight
months, while the same live weight is not obtained in the standard beef
breed until eighteen months.
These examples serve to illustrate the concept that, as an evolutionary
force, introgression is rather insignificant in natural biomes. It is almost al-
ways in the wake of human activity or the activities of their domesticated
animals that the process of introgression can and does result in new combi-
nations of gene pools from different species.
Edward N. Nelson
See also: Adaptive radiation; Biodiversity; Biogeography; Convergence

and divergence; Demographics; Extinctions and evolutionary explosions;
Gene flow; Genetic diversity; Genetic drift; Isolating mechanisms; Non-
random mating, genetic drift, and mutation; Population analysis; Popula-
tion fluctuations; Population genetics; Population growth; Punctuated
equilibrium vs. gradualism; Reproductive strategies; Speciation.

Anderson, Edgar. Introgressive Hybridization. New York: Hafner Press,
1968.
Arnold, M. L. Natural Hybridization and Evolution. New York: Oxford Uni-
versity Press, 1997.
Briggs, D., and S. M. Walters. Plant Variation and Evolution. New York: Cam-
84
Dobzhansky, Theodosius G. Genetics and the Origin of Species. 1951. 3d rev.

ed. Reprint. New York: Columbia University Press, 1982.
Endler, John A. Geographic Variation: Speciation and Clines. Princeton, N.J.:
Princeton University Press, 1977.
Grant, V. The Origin of Adaptations. New York: Columbia University Press,
1963.
Kimbel, William H., and Lawrence B. Martin, eds. Species, Species Concepts,
and Primate Evolution. New York: Plenum Press, 1993.
Ridley, M. Evolution. Boston: Blackwell Scientific Publications, 1993.
Roughgarden, Jonathan. Theory of Population Genetics and Evolutionary Ecol-
ogy: An Introduction. Upper Saddle River, N.J.: Prentice Hall, 1996.
Stebbins, G. L., Jr. Variation and Evolution in Plants. New York: Columbia
85
COEVOLUTION
Types of ecology: Community ecology; Evolutionary ecology
Coevolution is the interactive evolution of two or more species that results in a

mutualistic or antagonistic relationship.
W hen two or more different species evolve in a way that affects one
another’s evolution, coevolution is taking place. This interactive
type of evolution is characterized by the fact that the participant life-forms
are acting as a strong selective pressure upon one another over a period of
time. The coevolution of plants and animals, whether animals are consid-
ered strictly in their plant-eating role or also as pollinators, is abundantly
represented in every terrestrial ecosystem throughout the world where
flora has established itself. Moreover, the overall history of some of the
multitude of present and past plant and animal relationships is displayed
(although fragmentally) in the fossil record found in the earth’s crust.
Beginnings
The most common coevolutionary relationships between plants and ani-
mals concern plants as a food source. Microscopic, unicellular plants were
the earth’s first autotrophs (organisms that can produce their own organic
energy through photosynthesis, that is, from basic chemical ingredients
derived from the environment). In conjunction with the appearance of
autotrophs, microscopic, unicellular heterotrophs (organisms, such as ani-
mals, that must derive food from other sources, such as autotrophs) evolved
to exploit the autotrophs.
Sometime during the later part of the Mesozoic era, angiosperms, the
flowering plants, evolved and replaced most of the previously dominant
land plants, such as the gymnosperms and the ferns. New species of herbi-
vores evolved to exploit these new food sources. At some point, probably
during the Cretaceous period of the late Mesozoic era, animals became un-
intentional aids in the angiosperm pollination process. As this coevolution
proceeded, the first animal pollinators became more and more indispens-
able as partners to the plants.
Eventually, highly coevolved plants and animals developed relation-
ships of extreme interdependence, exemplified by the honeybees and their
coevolved flowers. This angiosperm-insect relationship is thought to have
arisen in the Mesozoic era by way of beetle predation, possibly on early,
magnolia-like angiosperms. The fossil record gives some support to this
86
Coevolution
theory. Whatever the exact route along which plant-animal pollination

partnerships coevolved, the end result was a number of plant and animal
species that gained mutual benefit from the new type of relationship.
Coevolutionary Relationships
Coevolved relationships include an immense number of relationships be-
tween plants and animals, and even between plants and other plants.
Among these coevolved situations can be found commensalisms, in which
different species have coevolved to live intimately with one another with-
out injury to any participant, and symbioses, in which species have co-
evolved to literally “live together.”
Such intertwined relationships can take the form of mutualism, in
which neither partner is harmed and indeed one or both benefit—as in
the relationships between fungi and algae in lichens, fungi and roots in
mycorrhizae, and ants and acacia trees in a symbiotic mutualism in which
the ants protect the acacias from herbivores. In parasitism, however, one
partner benefits at the expense of the other; a classic example is the rela-
tionship between the mistletoe parasite and the oak tree. Another coevolu-
A nonsymbiotic mutualism has coevolved between this hummingbird and the fun-
nel shape of the flower from which it extracts nectar with its long, pointed bill. The
plant has achieved greater evolutionary fitness through its ability to attract the
bird, which helps the plant propagate by facilitating pollination and seed dispersal.
At the same time, the hummingbird benefits from a source of nutrition tailored to
its anatomy and protected from competitors. (Corbis)
87
Coevolution
tionary relationship, predation, is restricted primarily to animal-animal re-

lationships (vertebrate carnivores eating other animals, most obviously),
although some plants, such as Venus’s flytrap, mimic predation in having
evolved means of trapping and ingesting insects as a source of food. Some
highly evolved fungi, such as the oyster mushroom, have evolved anesthe-
tizing compounds and other means of trapping protozoa, nematodes, and
other small animals.
One of the most obvious and complex coevolutionary relationships are
the mutualisms that have evolved between plants bearing fleshy fruits and
vertebrate animals, which serve to disperse the seeds in these fruits. Over
time, plants that produce these fruits have benefited from natural selection
because their seeds have enjoyed a high degree of survival and germina-
tion: Animals eat the fruits, whose seeds are passed through their digestive
systems (or regurgitated to feed offspring) unharmed; at times the seeds
are even encouraged toward germination as digestion helps break down
the seed coat. Furthermore, dispersal through the animals’ mobility allows
the seeds to enjoy more widely distributed propagation. The coevolution-
ary process works on the animals as well: Birds and animals that eat the
fruits enjoy a higher degree of survival, and so natural selection favors
both fleshy-fruit-producing plants and fleshy-fruit-eating animals. Similar
selection has favored the coevolution of flowers with colors and smells that
attract pollinators such as bees.
Eventually some plant-animal mutualisms became so intertwined that
one or both participants reached a point at which they could not exist with-
out the aid of the other. These obligatory mutualisms ultimately involve
other types of animal partners besides insects. Vertebrate partners such as
birds, reptiles, and mammals became involved in mutualisms with plants.
In the southwestern United States, for example, bats and the agave and sa-
guaro cactus have a special coevolutionary relationship: The bats, nectar
drinkers and pollen eaters, have evolved specialized feeding structures
such as erectile tongues similar to those found among moths and other in-
sects with similar lifestyles. In turn, angiosperms coevolutionarily in-
volved with bats have developed such specializations as bat-attractive
scents, flower structures that match the bats’ feeding habits and minimize
the chance of injuring the animals, and petal openings timed to the noctur-
nal activity of bats.
Defense Mechanisms
Coevolution is manifested in defense mechanisms as well as attractants:
Botanical structures and chemicals (secondary metabolites) have evolved
to discourage or to prevent the attention of plant eaters. These include the
88
Coevolution
development of spines, barbs, thorns, bristles, and hooks on plant leaves,

stems, and trunk surfaces. Cacti, hollies, and rose bushes illustrate this
form of plant strategy. Some plants produce chemical compounds that are
bitter to the taste or poisonous. Plants that contain organic tannins, such as
trees and shrubs, can partially inactivate animals’ digestive juices and cre-
ate cumulative toxic effects that have been correlated with cancer. Grasses
with a high silica content act to wear down the teeth of plant eaters. Ani-
mals have counteradapted to these defensive innovations by evolving a
higher degree of resistance to plant toxins or by developing more efficient
and tougher teeth with features such as harder enamel surfaces or the ca-
pacity of grinding with batteries of teeth.
Frederick M. Surowiec, updated by Christina J. Moose
See also: Adaptations and their mechanisms; Adaptive radiation; Coloni-

zation of the land; Convergence and divergence; Defense mechanisms; Evo-
lution: definition and theories; Evolution of plants and climates; Grazing
and overgrazing; Metabolites; Natural selection; Paleoecology; Pollina-
tion; Symbiosis.

Bakker, Robert T. The Dinosaur Heresies. Reprint. Secaucus, N.J.: Citadel,
2001.
Barth, Friedrich G. Insects and Flowers: The Biology of a Partnership. Trans-
lated by M. A. Biederman-Thorson. Princeton, N.J.: Princeton Univer-
sity Press, 1991.
Gilbert, Lawrence E., and Peter H. Raven, eds. Coevolution of Animals and
Plants. Austin: University of Texas Press, 1980.
Gould, Stephen Jay. The Panda’s Thumb. Reprint. New York: W. W. Norton,
1992.
Grant, Susan. Beauty and the Beast: The Coevolution of Plants and Animals.
New York: Charles Scribner’s Sons, 1984.
Hughes, Norman F. Paleobiology of Angiosperm Origins. Reprint. New York:
Cambridge University Press, 1994.
Thompson, John N. The Coevolutionary Process. Chicago: University of Chi-
cago Press, 1994.
89
COLONIZATION OF THE LAND
Types of ecology: Evolutionary ecology; Paleoecology
The advent of animals and plants on land during the Ordovician period added new
complexity to preexisting ecosystems and paved the way for land ecosystems. The
newly increased mass of vegetation on land served to stabilize soils against erosion
and promoted the weathering of their nutrient minerals. Arthropods, too, found a
place in this early ecosystem of nonvascular plants on land.
The appearance of animals and plants on land by the Middle Ordovi-

cian period, some 450 million years ago, was a major event in the evolution
of terrestrial ecosystems. Nevertheless, they probably were not the earth’s
first inhabitants; there is a fossil record of blue-green algae and other mi-
croscopic life well back into Precambrian time, as much as 3.5 billion years
ago. Indeed, it is doubtful that plants and animals visible to the naked eye
could have lived on land without preexisting microbial ecosystems, which
served to stabilize minerals in soils, to decompose and circulate organic
matter of dead organisms, and to oxygenate the atmosphere by photosyn-
thesis.
The increased mass of more complex animals and plants on land during
Ordovician time further stabilized soils, invigorated the recycling of or-
ganic matter, and boosted atmospheric oxygenation. In addition, large
plants provided greater depth and structure to terrestrial ecosystems than
was possible with microbes and so may have promoted photosynthetic ef-
ficiency, biological diversity, and perhaps also resistance to disturbance by
floods and storms. This self-reinforcing boost to terrestrial productivity
firmly established life on land.
Invasion from the Sea

Because there are marine fossils of plants and animals visible to the naked
eye in Precambrian rocks (at least 600 million years old), it has commonly
been assumed that the earliest creatures on land during the Ordovician
and Silurian periods invaded from the sea. Reasons advanced to explain
why the land was unavailable for marine creatures for more than 200 mil-
lion years include the lack of available oxygen, the poverty of terrestrial
microbial photosynthetic productivity, and an unpredictable land surface
of flash floods and erosional badlands. This view of an invasion from the
sea has been used to explain the origins of earliest land animals, which
probably were arthropods, such as millipedes and spiders. A tremendous
90
Colonization of the land
variety of fossil arthropods have been found in Cambrian, Ordovician, and

Silurian deposits of shallow seas and estuaries. Like modern marine crabs,
these creatures may have ventured out to a limited extent on land, and
some may have become more fully adapted to more difficult conditions
there. The external skeletons of arthropods, important for defense in the
sea, also are effective for support, movement, and preventing desiccation
on land.
On the other hand, millipedes and spiders are not very closely related
either to any known fossil or to living aquatic arthropods. A reassessment
of the earliest fossil scorpions, formerly regarded as possible early land an-
imals, has shown that they had a breathing apparatus that would have
been effective only in water. Substantial evolution on land must have oc-
curred to produce the earliest spiders and millipedes, perhaps from micro-
scopic early microbial feeders that have left no fossil record.
Immigrant vs. Indigenous Evolution

The idea of invasion of the land by marine and freshwater algae is sup-
ported to some extent by the close biochemical similarities between mod-
ern land plants and charophytes (a kind of pond weed commonly called
stonewort because of its calcified egg cells). Charophytes, however, are
very different from land plants, and it is unlikely that such soft-bodied
aquatic algae in the geological past were any more successful in colonizing
the land than are the mounds of rotting seaweed now thrown up on
beaches by storms.
Land plants differ from stoneworts and seaweeds in many ways: They
have a waxy and proteinaceous outer coating (cuticle) to prevent desicca-
tion and to allow the plant body to remain turgid through internal water
pressure; they have small openings (stomates) surrounded by cells that can
open and close the opening in order to control loss of water and oxygen
and intake of carbon dioxide; they have internal systems of support and
water transport, which include tubular thick-walled cells (hydroids) in
nonvascular plants, such as mosses and liverworts, and elongate cells with
helical or banded woody thickenings (tracheids) in vascular plants; they
have roots, unicellular root hairs, or rootlike organs (rhizoids) that gather
water and nutrients from soil; and they have propagules (spores) protected
from desiccation and abrasion by proteinaceous envelopes. To some bota-
nists, the coordinated evolution of all these features from aquatic algae is
extremely unlikely, notwithstanding the impressive diversity of algae to-
day. This consideration, plus the simple nature of the earliest fossil land
plants, has led to the argument that land plants evolved on land from mi-
croscopic algae already accustomed to such conditions.
91
The Earliest Land Ecosystems

While immigrant versus indigenous evolutionary origins of the earliest
land creatures remains a theoretical problem, there is fossil evidence of
very early land ecosystems. In Late Ordovician rocks are found the earliest
spores of land plants. Most of them are smooth and closely appressed in
groups of four, somewhat similar to spores of liverworts and mosses today.
This is not to say that they belonged to liverworts and mosses; no clear fos-
sils of land plants visible to the naked eye have yet been found in rocks of
this age. Early moss and liverwort ancestors are found in Silurian rocks,
but so are extinct nonvascular plants, such as nematophytes. These early
experiments in the evolution of land plants had tissues supported by
densely interwoven proteinaceous tubes. In life, they had the rubbery tex-
ture of a mushroom and a variety of bladelike and elongate forms similar
to those of some living algae.
Although the botanical affinities of the earliest spores of nonvascular
land plants remain unclear, there is evidence that they grew in clumps.
Buried soils of Late Ordovician age have been found with surficial erosion
scours of the kind formed by wind around clumps of vegetation. The
clumps are represented by gray spots from the reducing effect of remnant
organic matter. Burrows also have been found in Late Ordovician buried
soils as an indication of animals in these early land ecosystems. The fossil
burrows are quite large (2 to 21 millimeters). They are similar in their
clayey linings, backfill structures, and fecal pellets to the burrows of mod-
ern roundback millipedes. The buried soils are calcareous and strongly
ferruginized—indications that they were nutrient-rich, periodically dry,
and well drained, as are modern soils preferred by millipedes. Actual fos-
sils of millipedes have not yet been found in rocks older than Late Silurian,
so all that can be said at present is that these very early animals on land
were in some ways like millipedes.
Diversification of Life on Land

By Silurian time (some 438 million years ago) there was a considerable di-
versification of life on land. Spores of fungi and of vascular land plants
have been found fossilized in Early Silurian rocks. During Mid-Silurian
time, there were small, leafless plants with bifurcating rhizomes and pho-
tosynthetic stems terminated by globular, spore-bearing organs. These
matchstick-sized fossil plants have been called Cooksonia. Although not
so well preserved as to show their water-conducting cells, they have been
regarded as the earliest representatives of the extinct group of vascu-
lar plants called rhyniophytes. In Devonian rocks (some 408 million
years old), some well-preserved rhyniophytes are known to have been
92
true vascular plants, but there are other plants similar in general appear-
ance that had simpler thick-walled conducting cells like those of non-
vascular plants. By Devonian time, there were also vascular plants with
spore-bearing organs borne above lateral branches (zosterophylls), plants
with true roots and spore-bearing organs borne in clusters (trimero-
phytes), and spore-producing plants with woody roots and tree trunks
(progymnosperms). The evolution of the earliest vascular plant cover
on land, and of the first forests, involved different kinds of plants now ex-
tinct.
To fossil millipedes of Silurian age were added during Devonian time
spiders, centipedes, springtails (Collembola), and bristletails (Thysanura).
The earliest vertebrates on land are known from bones of extinct amphibi-
ans (Ichthyostegalia) and from footprints of Devonian age, some 370 million
years old.
This great Silurian and Devonian evolutionary radiation promoted en-
vironmental changes similar to those initiated by the first colonization of
land by plants and animals, as well as some new changes. For example, the
formation of charcoal from wildfires in woodlands and the accumulation
of peat in swamps were ways of burying carbon that otherwise might have
decayed or digested into carbon dioxide in the atmosphere. Removal of
carbon dioxide in this way allowed increased oxygenation of the atmo-
sphere. Oxygenation was kept within bounds by increased flammability of
woodlands when oxygen reached amounts much in excess of the present
atmospheric level.
Late Devonian ecosystems were very different from modern ones. Ma-
jor ecological roles, such as insect-eating large animals on land, were still
being added. More changes were to come, but the world at that time would
have seemed a much more familiar place than the meadows of Cooksonia
during the Silurian, the patchy cover of Ordovician nonvascular plants,
and the red and green microbial earths of earlier times.
Gregory J. Retallack
See also: Coevolution; Convergence and divergence; Evolution: definition

and theories; Evolution: history; Evolution of plants and climates; Extinc-
tions and evolutionary explosions; Mycorrhizae; Natural selection; Paleo-
ecology; Punctuated equilibrium vs. gradualism.

Gordon, M. S., and E. C. Olson. Invasions of the Land: The Transition of Organ-
isms from the Aquatic to Terrestrial Life. New York: Columbia University
Press, 1995.
93
Little, C. The Colonization of the Land. Cambridge, England: Cambridge Uni-

Schopf, J. William, ed. Major Events in the History of Life. Boston: Jones and
Bartlett, 1992.
Schumm, Stanley A. The Fluvial System. New York: Wiley-Interscience,
1977.
Stanley, Steven M. Earth and Life Through Time. 2d ed. New York: W. H. Free-
man, 1989.
Stebbins, G. L., and G. J. C. Hill. “Did Multicellular Plants Invade the
Land?” American Naturalist 115 (1980): 342-353.
Wright, V. P., and Alfred Fischer, eds. Paleosols: Their Recognition and Inter-
pretation. Princeton, N.J.: Princeton University Press, 1986.
Zimmer, Carl. At the Water’s Edge: Macroevolution and the Transformation of
Life. New York: Free Press, 1998.
94
COMMUNICATION
Types of ecology: Behavioral ecology; Chemical ecology
In animal communication, information is exchanged through signals. Such sig-

nals are vital for survival, finding mates, and rearing young.
A simple definition of animal communication is the transmission of in-

formation between animals by means of signals. Developing a more
precise definition is difficult because of the broad array of behaviors that
are considered messages or signals and the variety of contexts in which
these behaviors may occur. Animal signals can be chemical, visual, audi-
tory, tactile, or electrical. The primary means of communication used
within a species will depend upon its sensory capacities and its ecology.
Pheromones
Of the modes of communication available, chemical signals, or phero-
mones, are assumed to have been the earliest signals used by animals.
Transmission of chemical signals is not affected by darkness or by obsta-
cles. One special advantage is that the sender of a chemical message can
leave the message behind when it moves. The persistence of the signal may
also be a disadvantage when it interferes with transmission of newer infor-
mation. Another disadvantage is that the transmission is relatively slow.
The speed at which a chemical message affects the recipient varies. Some
messages have an immediate effect on the behavior of recipients. Alarm and
sex-attractant pheromones of many insects, aggregation pheromones in
cockroaches, or trail substances in ants are examples. Other chemical mes-
sages, primers, affect recipients more slowly, through changes in their physi-
ology. Examples of primers include pheromones that control social structure
in hive insects such as termites. Reproductive members of the colony secrete
a substance that inhibits the development of reproductive capacity in other
hive members. The chemicals important for controlling the hive are spread
through grooming and food sharing (trophallaxis). Chemical communica-
tion is important not only among social and semisocial insects but also
among animals, both vertebrate and invertebrate. Particularly common is
the use of a pheromone to indicate that an animal is sexually receptive.
Visual Signals
Visual communication holds forth the advantage of immediate transmis-
sion. A visual signal or display is also able to encode a large amount of in-
95
Communication
Image Not Available
formation, including the location of the sender. Postures and movements

of parts of an animal’s body are typical elements of visual communication.
Color and timing are additional means of providing information. Some vi-
sual signals are discrete; that is, the signal shows no significant variation
from performance to performance. Other displays are graded so that the
information content of the signal can be varied. An example of a graded
display is found in many of the threat or aggressive postures of birds.
Threat postures of the chaffinch vary between low-intensity and high-
intensity postures. The elevation of the crest varies in ways that indicate
the bird’s relative readiness for combat. The song spreads of red-winged
blackbirds and cowbirds show variation in intensity. In red-winged black-
birds, the red epaulets, or shoulder patches, are exposed to heighten the
effect of the display. Discrete and graded signals may be used together to
increase the information provided by the signal. In zebras, ears back indi-
cates a threat and ears up indicates greeting. The intensity of either mes-
sage is shown by the degree to which the mouth is held open. A widely
open mouth indicates a heightened greeting or threat.
Visual displays depend upon the presence of light or the production of
light. The ability to produce light, bioluminescence, is found most fre-
quently in aquatic organisms, but its use in communication is probably
best documented in fireflies, beetles belonging to the family Lampyridae.
96
Communication
Firefly males advertise their presence by producing flashes of light in a

species-specific pattern. Females respond with simple flashes, precisely
timed, to indicate that they belong to the appropriate species. This commu-
nication system is used to advantage by females in a few predatory species
of the genus Photuris. After females of predatory species have mated with
males of their own species, they attract males of other species by mimick-
ing the responses of the appropriate females. The males that are tricked are
promptly eaten. The luminescence of fireflies does not attract a wide vari-
ety of nocturnal predators, because their bodies contain a chemical that
makes them unpalatable.
Visual displays are limited in the distance over which they can be used
and are easily blocked by obstacles. Visual communication is important in
primates, birds, and some insects, but can be dispensed with by many spe-
cies that do not have the necessary sensory capacities.
Auditory Communication
The limitation of visual communication is frequently offset by the coupling
of visual displays with other modes of communication. Visual displays can
be coupled with auditory communication, for example. There are many ad-
vantages to using sound: It can be used in the dark, and it can go around
obstacles and provide directional information. Because pitch, volume, and
temporal patterns of sound can be varied, extremely complex messages
can be communicated. The auditory communication of many bird species
has been studied intensively. Bird vocalizations are usually classified into
two groups, calls and songs. Calls are usually brief sounds, whereas songs
are longer, more complex, and often more suited to transmission over dis-
tances.
The call repertoire of a species serves a broad array of functions. Many
young birds use both a visual signal, gaping, and calling in their food beg-
ging. Individuals that call more may receive more food. Begging calls and
postures may also be used by females in some species to solicit food from
mates. One call type that has been intensively studied is the alarm call.
Alarm calls of many species are similar, and response is frequently interspe-
cific (that is, interpretable by more than one species). Alarm calls are likely
to be difficult to locate, a definite advantage to the individual giving the
call. Calls used to gather individuals for mobbing predators are also similar
in different species. Unlike alarm calls, mobbing calls provide good direc-
tional information, so that recruitment to the mobbing effort can be rapid.
Call repertoires serve birds in a great variety of contexts important for
survival of the individual. Song, on the other hand, most often serves a re-
productive function, that of helping a male hold a territory and attract a
97
Communication
mate. Songs are species-specific, like the distinctive markings of a species.

In some cases, songs are more distinctive than physical appearance. The
chiffchaff and willow warbler were not recognized as separate species un-
til an English naturalist named Gilbert White discovered, by examining
their distinctive songs, that they are separate. The North American wood
and hermit thrushes can also be distinguished more readily by song than
by appearance. Birdsong can communicate not only the species of the indi-
vidual singing but also information about motivational state. Most singing
is done by males during the breeding season. In many species, only the
male sings. In some species, females sing as well. Their songs may be simi-
lar to the songs of the males of their species or they may be distinctive. If
the songs are similar to those of the males, the female may sing songs infre-
quently and with less volume. In some instances, the female song serves to
notify her mate of her location. An interesting phenomenon found in some
species is duetting, in which the male and female develop a duet. Mates
may sing in alternate and perfectly timed phrases, as is done by the African
boubou shrike, Laniarius aethiopicus. An individual shrike can recall its
mate by singing the entire song alone.
Individuals in some bird species have a single song, and individuals of
other species have repertoires of songs. Average repertoire size of the indi-
vidual is characteristic of a species. Whether songs in repertoires are
shared with neighbors or unique to the individual is also characteristic of a
species. Sharing songs with neighbors permits song matching in counter-
singing. Cardinals and tufted titmice are species that frequently match
songs in countersinging. Possible uses for matching are facilitating the rec-
ognition of intruders and indicating which neighbor has the attention of a
singer. Some species of birds have dialects. The species-specific songs of
one geographic region can be differentiated from the song of another geo-
graphic region. The development of dialects may be useful in maintaining
local adaptations within a species, provided that females select mates of
the same dialect as their fathers.
Although auditory signals of birds have received a disproportionate
share of attention in the study of animal communication, auditory com-
munication is used by a broad spectrum of animals. Crickets have species-
specific songs to attract females and courtship songs to encourage an ap-
proaching female. The ears of most insects can hear only one pitch, so the
temporal pattern of sound pulses is the feature by which a species can be
identified. Vervet monkeys use three different alarm calls, depending
upon the kind of threat present; they respond to the calls appropriately by
looking up, looking down, or climbing a tree, depending upon the kind of
call given.
98
Communication
Tactile Communication
Tactile communication differs significantly from other forms of communi-
cation in that it cannot occur over a distance. This form of communication
is important in many insects, equipped as they are with antennae rich in re-
ceptors. Shortly after a termite molts, for example, it strokes the end of the
abdomen of another individual with its antennae and mouthparts. The in-
dividual receiving this signal responds by extruding a fluid from its
hindgut. Tactile signals are frequently used in eliciting trophallaxis (food
sharing) in social insects. Tactile signals are also important in the copula-
tory activity of a number of vertebrates.
Additional channels of communication available in animals are electrical
and surface vibration. Many modes of communication are used in combina-
tion with other modes. The channels used will depend in part on the sensory
equipment of the species, its ecology, and the particular context. Most mes-
sages will be important either for the survival of the individual or the group
or for the individual’s ability to transmit its genes to the next generation.
See also: Altruism; Defense mechanisms; Displays; Ethology; Hierarchies;

Insect societies; Mammalian social systems; Mimicry; Pheromones; Poi-
sonous animals; Predation; Reproductive strategies; Territoriality and ag-
gression.

De Waal, Frans. Chimpanzee Politics: Power and Sex Among Apes. New York:
Harper & Row, 1982.
Goodall, Jane. In the Shadow of Man. Rev. ed. Boston: Houghton Mifflin,
2000.
Gould, James L. Ethology. New York: W. W. Norton, 1982.
Grier, James W. Biology of Animal Behavior. 2d ed. St. Louis: Times Mirror/
Mosby, 1992.
Hart, Stephen. The Language of Animals. New York: H. Holt, 1996.
Hauser, Marc D. The Evolution of Communication. Cambridge, Mass.: MIT
Press, 1996.
Peters, Roger. Mammalian Communication: A Behavioral Analysis of Meaning.
Monterey, Calif.: Brooks/Cole, 1980.
Roitblat, Herbert L., Louis M. Herman, and Paul E. Nachtigall, eds. Lan-
guage and Communication: Comparative Perspectives. Hillsdale, N.J.: Law-
rence Erlbaum, 1993.
Wilson, Edward O. Insect Societies. Cambridge, Mass.: Harvard University
Press, 1971.
99
COMMUNITIES: ECOSYSTEM
INTERACTIONS
Types of ecology: Community ecology; Ecosystem ecology
Ecosystems are complex organizations of living and nonliving components. They

are frequently named for their dominant biotic or physical features (such as marine
kelp beds or coniferous forests). Communities are groups of species usually classi-
fied according to their most prominent members (such as grassland communities
or shrub communities). The interactions between species and their ecosystems
have lasting impacts on both.
I n an ecological sense, a community consists of all populations residing

in a particular area. Examples of communities range in scale from all the
trees in a given watershed, all soil microbes on an agricultural plot, or all
phytoplankton in a particular harbor to all plants, animals, and microbes
in vast areas, such as the Amazon River basin or the Chesapeake Bay.
An ecosystem consists of the community of species as well as the envi-
ronment of a given site. A forest ecosystem would include all living plants
and animals, along with climate, soils, disturbance, and other abiotic (non-
living) factors. An estuarine ecosystem, likewise, would include all the liv-
ing things present in addition to climate, currents, salinity, nutrients, and
more.
Interactions between species in communities and ecosystems range
from mutually beneficial to mutually harmful. One such category of inter-
action is mutualism, which usually involves two species. Both species de-
rive benefit from a mutualism. Commensalism is used to describe a situa-
tion in which one species benefits without harming the other. If the two
species are neither helped nor harmed, a neutralism is said to occur, and an
amensalism happens when one species is harmed while the other remains
unaffected. During competition, both species involved are negatively af-
fected. A number of terms are used to describe a relationship in which one
species benefits at another’s expense, including herbivory, predation, par-
asitism, and pathogenicity. The choice of term more often than not de-
pends on the relative sizes of the species involved.
Competition
Plants typically compete for resources, such as light, space, nutrients, and
water. One way an individual may outcompete its neighbors is to outgrow
100
Communities: ecosystem interactions
them, thus capturing more sunlight for itself (and thus producing more
sugars and other organic molecules for itself). Another way is to be more
fecund than the neighbors, flooding the surroundings with one’s progeny
and thereby being more likely to occupy favorable sites for reproduction.
For example, in closed forests treefall gaps are quickly filled with growth
from the canopy, thus shading the ground and making it more difficult for
competing seedlings and saplings to survive.
Plants compete in the root zone as well, as plants with a more extensive
root network can acquire more of the water and other inorganic nutrients
necessary for growth and reproduction than can their competitors. In
semiarid areas, for example, trees often have trouble colonizing grasslands
because the extensive root systems of grasses are much more effective in
capturing available rainwater.
Sometimes plants resort to chemical “warfare,” known as allelopathy,
in order to inhibit the growth of competitors in the surrounding area. The
existence of allelopathy remains a controversial topic, and simpler expla-
nations have been offered for many previously alleged instances of the
phenomenon. Allelopathy cannot be rejected outright; however, the con-
troversy most likely proves only that many aspects of nature cannot be pi-
geonholed into narrow explanations.
Competition involves a cost in resources devoted to outgrowing or
outreproducing the neighbors. Because of the cost, closely related, compet-
ing species will diverge in their ecological requirements. This principle is
known as competitive exclusion.
Mutualism, Commensalism, and Parasitism

Many flowering plants could not exist without one of the most important
mutualisms of all: pollination. In concept, pollination is simple: In ex-
change for carrying out the physical work of exchanging genetic material
(in pollen form) between individual plants (thus enabling sexual repro-
duction), the carrier is rewarded with nutrients in the form of nectar or
other materials. Many types of animals are involved in pollination: insects
such as bees, flies, and beetles; birds, particularly the hummingbirds; and
mammals such as bats.
Another highly important mutualism is that between plant roots and
fungal hyphae, or mycorrhizae. Mycorrhizae protect plant roots from
pathogenic fungi and bacteria; their most important role, however, is to en-
hance water and nutrient uptake by the plant. In fact, regeneration of some
plants is impossible in the absence of appropriate mycorrhizae. Mycor-
rhizae benefit, in turn, by receiving nutrients and other materials synthe-
sized by the host plant. There are two types of mycorrhizae: ectomycor-
101
rhizae, whose hyphae may fill the space between plant roots but do not
penetrate the roots themselves; and vesicular-arbuscular mycorrhizae,
whose hyphae penetrate and develop within root cells.
Few people can envision a swamp in the southeastern United States
without thinking of bald cypress trees (Taxodium distichum) draped in ethe-
real nets of Spanish moss (Tillandsia usneoides), which is actually not a moss
but a flowering plant in the monocot family Bromeliaceae. Tillandsia is an
epiphyte, a plant that grows on the stems and branches of a tree. Epi-
phytism is one of the most common examples of a commensalism, in
which one organism, for instance the epiphyte, benefits without any de-
monstrable cost to the other, in this case the host tree. Epiphytes are com-
mon in tropical rain forests and include orchids, bromeliads, cacti, and
ferns. In temperate regions more primitive plants, such as lichens, are more
likely to become epiphytes.
Not all epiphytes are commensal, however. In the tropics, strangler figs,
such as Ficus or Clusia, begin life as epiphytes but send down roots that in
time completely encircle and kill the host. Mistletoes, such as Phoradendron
or Arceuthobium, may draw off the photosynthetic production of the host,
thus severely depleting its resources.
Herbivory and Pathogenicity

Plants, because of their ability to harvest light energy from the sun to pro-
duce the organic nutrients and building blocks necessary for life, are the
primary producers of most of the earth’s ecosystems. Thus, they face an
onslaught of macroscopic and microscopic consumers. If macroscopic, the
consumers are generally regarded as herbivores (plant-eating animals); if
microscopic, they are pathogens. Either way, herbivores and pathogens
generally devour the tissues of the host.
Plants have evolved a number of defense mechanisms in response to
pressure from herbivores and pathogens. Some responses may be mechan-
ical. For example, trees on an African savanna may evolve greater height to
escape grazing pressures from large herbivores, but some large herbivores,
specifically giraffes, may evolve to grow to greater heights as well. Plants
may encase themselves in nearly indestructible outer coatings or arm
themselves with spines in order to discourage grazers.
Other responses may be chemical. Cellulose, one of the important
chemical components of plant tissues such as wood, is virtually indigest-
ible—unless the herbivore itself hosts a bacterial symbiont in its stomach
that can manage the job of breaking down cellulose. Other chemicals, such
as phenols and tannins—the class of compounds that gives tea its brown
color—are likewise indigestible, thus discouraging feeding by insects.
102
Plants produce a wide range of toxins, such as alkaloids, which poison or

kill herbivores. A number of hallucinogenic drugs are made from plant al-
kaloids.
Phytoalexins are another group of defensive compounds produced by
plants in response to bacterial and fungal pathogens. Substances present in
the cell walls of bacteria and fungi are released via the action of plant en-
zymes and spread throughout the plant. The bacterial and fungal sub-
stances function as hormones to stimulate, or elicit, phytoalexin produc-
tion. Hence, these substances are referred to as elicitors. The phytoalexins
act as antibiotics, killing the infective agents. Tannins, phenols, and other
compounds also serve to defend against pathogen attack.
David M. Lawrence
See also: Allelopathy; Animal-plant interactions; Biodiversity; Biogeog-

raphy; Biological invasions; Coevolution; Communities: structure; Compe-
tition; Defense mechanisms; Food chains and webs; Lichens; Metabolites;
Mycorrhizae; Pollination; Predation; Speciation; Succession; Symbiosis;

Barbour, Michael G., et al. Terrestrial Plant Ecology. 3d ed. Menlo Park,
Calif.: Benjamin Cummings, 1999.
Barnes, Burton V., Donald R. Zak, Shirley R. Denton, and Stephen H. Spurr.
Forest Ecology. 4th ed. New York: John Wiley & Sons, 1998.
Mitch, William J., and James G. Gosselink. Wetlands. 3d ed. New York: John
Wiley & Sons, 2000.
Morin, Peter J. Community Ecology. Oxford, England: Blackwell Science,
1999.
103
COMMUNITIES: STRUCTURE
Type of ecology: Community ecology
An ecological community is the assemblage of species found in a given time and

place. The species composition of different ecosystems and the ways in which they
maintain equilibrium and react to disturbances are manifestations of the commu-
nity’s stability.
T he populations that form a community interact through the processes

of competition, predation, parasitism, and mutualism. The structures
of communities are determined, in part, by the nature and strength of these
biotic factors. Abiotic factors (physical factors such as temperature, rain-
fall, and soil fertility) are the other set of important influences determining
community structure. An ecological community together with its physical
environment is called an ecosystem. No ecosystem can be properly under-
stood without a careful study of the biotic and abiotic factors that shape it.
Energy Flow
The most common way to characterize a community functionally is by de-
scribing the flow of energy through it. Based on the dynamics of energy
flow, organisms can be classified into three groups: those that obtain en-
ergy through photosynthesis (called producers), those that obtain energy
by consuming other organisms (consumers), and those that decompose
dead organisms (decomposers). The pathway through which energy trav-
els from producer through one or more consumers and finally to de-
composer is called a food chain. Each link in a food chain is called a trophic
level. Interconnected food chains in a community constitute a food web.
Very few communities are so simple that they can be readily described
by a food web. Most communities are compartmentalized: A given set of
producers tends to be consumed by a limited number of consumers, which
in turn are preyed upon by a smaller number of predators, and so on. Al-
ternatively, consumers may obtain energy by specializing on one part of
their prey (for example, some birds may eat only seeds of plants) but utilize
a wide range of prey species. Compartmentalization is an important fea-
ture of community structure; it influences the formation, organization, and
persistence of a community.
Dominant and Keystone Species

Some species, called dominant species, can exert powerful control over the
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Communities: structure
abundance of other species because of the dominant species’ large size, ex-
tended life span, or ability to monopolize energy or other resources. Com-
munities are named according to their dominant species: for example, oak-
hickory forest, redwood forest, sagebrush desert, and tall-grass prairie.
Some species, called keystone species, have a disproportionately large ef-
fect on community structure. These interact with other members of the
community in such a way that loss of the keystone species can lead to the
loss of many other species. Keystone species may also be the dominate spe-
cies, but they may also appear insignificant to the community until they
are gone. For example, cordgrass (Spartina) is the dominant plant in many
tidal estuaries, and it is also a keystone species because so many members
of the community depend on it for food and shelter.
The species that make up a community are seldom distributed uni-
formly across the landscape; rather, some degree of patchiness is character-
istic of virtually all species. There has been conflicting evidence as to the
nature of this patchiness. Moving across an environmental gradient (for
example, from wet to dry conditions or from low to high elevations), there
is a corresponding change in species and community composition. Some
studies have suggested that changes in species composition usually occur
along relatively sharp boundaries and that these boundaries mark the bor-
der between adjacent communities. Other studies have indicated that spe-
cies tend to respond individually to environmental gradients and that
community boundaries are not sharply defined; rather, most communities
broadly intergrade into one another, forming what is often called an
ecotone.
Degrees of Species Interaction

These conflicting results have fueled a continuing debate as to the underly-
ing nature of communities. Some communities seem to behave in a coordi-
nated manner. For example, if a prairie is consumed by fire, it regenerates
in a predictable sequence, ultimately returning to the same structure and
composition it had before the fire. This process, called ecological succes-
sion, is to be expected if the species in a community have evolved together
with one another. In this case, the community is behaving like an organism,
maintaining its structure and function in the face of environmental distur-
bances and fluctuations (as long as the disturbances and fluctuations are
not too extreme). The existence of relatively sharp boundaries between ad-
jacent communities supports this explanation of the nature of the commu-
nity.
In other communities, it appears that the response to environmental
fluctuation or disturbance is determined by the evolved adaptations of the
105
species available. There is no coordinated community response but rather

a coincidental assembly of community structure over time. Some sets of
species interact together so strongly that they enter a community together,
but there is no evidence of an evolved community tendency to resist or ac-
commodate environmental change. In this case, the community is formed
primarily of species that happen to share similar environmental require-
ments.
Competition and Predation

Disagreement as to the underlying nature of communities usually reflects
disagreement about the relative importance of the underlying mechanisms
that determine community structure. Interspecific competition has long
been invoked as the primary agent structuring communities. Competition
is certainly important in some communities, but there is insufficient evi-
dence to indicate how widespread and important it is in determining com-
munity structure. Much of the difficulty occurs because ecologists must in-
fer the existence of past competition from present patterns in communities.
It appears that competition has been important in many vertebrate com-
munities and in communities dominated by sessile organisms, such as
plants. It does not appear to have been important in structuring communi-
ties of plant-eating insects. Furthermore, the effects of competition typi-
cally affect individuals that use identical resources, so that only a small
percentage of species in a community may be experiencing significant
competition at any time.
The effects of predation on community structure depend on the nature
of the predation. Keystone predators usually exert their influence by prey-
ing on species that are competitively dominant, thus giving less competi-
tive species a chance. Predators that do not specialize on one or a few spe-
cies may also have a major effect on community structure, if they attack
prey in proportion to their abundance. This frequency-dependent preda-
tion prevents any prey species from achieving dominance. If a predator is
too efficient, it can drive its prey to extinction, which may cause a selective
predator to become extinct as well. Predation appears to be most important
in determining community structure in environments that are predictable
or unchanging.
Disasters and Catastrophes

Chance events can also influence the structure of a community. No envi-
ronment is completely uniform. Seasonal or longer-term environmental
fluctuations affect community structure by limiting opportunities for colo-
nization, by causing direct mortality, or by hindering or exacerbating the
106
The checkerboard pattern of clear-cutting in forests of the Pacific Northwest

threatens the survival of the northern spotted owl, the marbled murrelet, Vaux’s
swift, and the red tree vole, even though fragments of the community remain. Such
disruptions of community structure can be mitigated by thinning to sustain
mixed-age, mixed-species trees. (PhotoDisc)
effects of competition and predation. Furthermore, all communities expe-

rience at least occasional disturbance: unpredictable, seemingly random
environmental changes that may be quite severe.
It is useful in this regard to distinguish between regular disturbances
and rarer, more frequent catastrophic events. For example, fire occurs so
often in tall-grass prairies that most of the plant species have become fire-
adapted—they have become efficient at acquiring nutrients left in the ash
and at sprouting or germinating quickly after a fire. In contrast, the 1980
eruption of Mount St. Helens, a volcanic peak in Washington State, was so
violent and so unexpected that no members of the nearby community were
adequately adapted to such conditions.
Natural disturbances occur at a variety of scales. Small-scale distur-
bances may simply create small openings in a community. In forests, for
example, wind, lightning, and fungi cause single mature trees to die and
fall, creating gaps that are typically colonized by species requiring such
openings. Large disturbances are qualitatively different from small distur-
bances in that large portions of a community may be destroyed, including
some of the ability to recover from the disturbance. For example, following
107
a large, intense forest fire, some tree species may not return for decades or
centuries, because their seeds were consumed by the fire, and colonizers
must travel a long distance.
Early ecologists almost always saw disturbances as destructive and dis-
ruptive for communities. Under this assumption, most mathematical mod-
els portrayed communities as generally being in some stable state; if a dis-
turbance occurred, the community inevitably returned to the same (or
some alternative) equilibrium. It later became clear, however, that natural
disturbance is a part of almost all natural communities. Ecologists now rec-
ognize that few communities exhibit an equilibrium; instead, communities
are dynamic, always responding to the last disturbance.
Long-Term Community Dynamics

The evidence suggests that three conclusions can be drawn about the long-
term dynamics of communities. First, it can no longer be assumed that all
communities remain at equilibrium until changed by outside forces. Dis-
turbances are so common, at so many different scales and frequencies, that
the community must be viewed as an entity that is constantly changing as
its constituent species readjust to disturbance and to one another.
Second, communities respond in different ways to disturbance. A com-
munity may exhibit resistance, not markedly changing when disturbance
occurs, until it reaches a threshold and suddenly and rapidly shifts to a
new state. Alternatively, a community may exhibit resilience by quickly re-
turning to its former state after a disturbance. Resilience may occur over a
wide range of conditions and scales of disturbance in a dynamically robust
system. On the other hand, a community that exhibits resilience only
within a narrow range of conditions is said to be dynamically fragile.
Finally, there is no simple way to predict the stability of a community.
At the end of the 1970’s, many ecologists predicted that complex commu-
nities would be more stable than simple communities. It appeared that sta-
bility was conferred by more intricate food webs, greater structural com-
plexity, and greater species richness. On the basis of numerous field
studies and theoretical models, many ecologists now conclude that no
such relationship exists. Both very complex communities, such as tropical
rain forests, and very simple communities, such as Arctic tundra, may be
very fragile.
Studying Communities
Most communities consist of thousands of species, and their complexity
makes them very difficult to study. Most community ecologists specialize
in taxonomically restricted subsets of communities (such as plant commu-
108
nities, bird communities, insect communities, or moss communities) or in

functionally restricted subsets of communities (such as soil communities,
tree-hole communities, pond communities, or detrivore communities).
The type of community under investigation and the questions of inter-
est determine the appropriate methods of study. The central questions in
most community studies are how many species are present and what is the
abundance of each. The answers to these questions can be estimated using
mark-recapture methods or any other enumeration method.
Often the aim is to compare communities (or to compare the same com-
munity at different times). A specialized parameter called similarity is
used to compare and classify communities; more than two dozen measures
of similarity are available. Measures of similarity are typically subjected to
cluster analysis, a set of techniques that groups communities on the basis
of their similarity.
Many multivariate techniques are used to search for patterns in com-
munity data. Direct gradient analysis is the simplest of these techniques; it
is used to study the distribution of species along an environmental gradi-
ent. Ordination includes several methods for collapsing community data
for many species in many communities along several environmental gradi-
ents onto a single graph that summarizes their relationships and patterns.
Community Disturbance
At the most basic level, destruction of a community eliminates the species
that make up the community. If the community is restricted in its extent,
and if its constituent species are found nowhere else, those species become
extinct. If the community covers a large area or is found in several areas, lo-
cal extinction of species may occur without causing global extinction.
Destruction of a community can cause unexpected changes in environ-
mental conditions that were modified by the intact community. Even par-
tial destruction of an extensive community can eliminate species. For ex-
ample, the checkerboard pattern of clear-cutting in Douglas fir forests of
the Pacific Northwest threatens the survival of the northern spotted owl,
the marbled murrelet, Vaux’s swift, and the red tree vole, even though
fragments of the community remain. Many fragments are simply too small
to support these species. A Douglas fir forest is regenerated following cut-
ting, but this young, even-aged stand is so different from an old, mixed-age
forest that it functions as a different type of community.
Altering the population of one species can affect others in a community.
The black-footed ferret was once found widely throughout central North
America as a predator of prairie dogs. As prairie dogs were poisoned,
drowned, and shot throughout their range, the number of black-footed fer-
109
rets declined. The species nearly became extinct, and an attempt to in-
crease their numbers and preserve the species was instituted in the late
1980’s in a Wyoming breeding program.
Introducing a new species into a community can severely alter the inter-
actions in the community. The introduction of the European rabbit into
Australia led to a population explosion of rabbits, excessive predation on
vegetation, and resulting declines in many native marsupials.
Finally, it appears that many communities exhibit stability thresholds; if
a community is disturbed beyond its threshold, its structure is perma-
nently changed. For example, acid deposition in lakes is initially buffered
by natural processes. As acid deposition exceeds the buffering capacity of a
lake, it causes insoluble aluminum in the lake bottom to become soluble,
and this soluble aluminum kills aquatic organisms directly or by making
them more susceptible to disease. The lesson is clear: It is far easier to dis-
rupt or destroy natural systems (even accidentally) than it is to restore or
reconstruct them.
Alan D. Copsey, updated by Bryan Ness
See also: Animal-plant interactions; Biodiversity; Biogeography; Biologi-

cal invasions; Coevolution; Communities: ecosystem interactions; Compe-
tition; Defense mechanisms; Eutrophication; Food chains and webs; Inva-
sive plants; Predation; Speciation; Species loss; Succession; Trophic levels

Aber, John D., and Jerry M. Melillo. Terrestrial Ecosystems. 2d ed. San Diego:
Harcourt, 2001.
Begon, Michael, John L. Harper, and Colin R. Townsend. Ecology: Individ-
uals, Populations, and Communities. 3d ed. Cambridge, Mass.: Blackwell
Science, 1996.
Bormann, Frank H., and Gene E. Likens. “Catastrophic Disturbance and
the Steady State in Northern Hardwood Forests.” American Scientist 67
(1979): 660-669.
Goldammer, J. G., ed. Tropical Forests in Transition: Ecology of Natural and
Anthropogenic Disturbance Processes. Boston: Springer-Verlag, 1992.
Krebs, Charles J. Ecology: The Experimental Analysis of Distribution and Abun-
dance. 5th ed. San Francisco: Benjamin Cummings, 2001.
Pickett, S. T. A., and P. S. White, eds. The Ecology of Natural Disturbance and
Patch Dynamics. Orlando, Fla.: Academic Press, 1985.
Pielou, E. C. The Interpretation of Ecological Data: A Primer on Classification
and Ordination. New York: John Wiley & Sons, 1984.
110
COMPETITION
Types of ecology: Behavioral ecology; Community ecology
Competition is the conflict between different organisms for control of food, natural
resources, territories, mates, and other aspects of survival. Competition can occur
between individuals of the same species or between individuals of different species.
In either case, it is natural selection for the fittest organisms and species; therefore,
it is a major driving force in evolution.
T he science of ecology can best be defined as the experimental analysis

of the distribution and abundance of organisms. Natural selection in-
fluences the distribution and abundance of organisms from place to place.
The possible selecting factors include physical factors (temperature and
light, for example), chemical factors such as water and salt, and species in-
teractions. Any of these factors can influence the survivability of organ-
isms in any particular environment. According to ecologist Charles Krebs,
species interactions include four principal types: mutualism, which is the
living together of two species that benefit each other (for example, humans
and their pets); commensalism, which is the living together of two species
that results in a distinct benefit (or number of benefits) to one species while
the other remains unhurt (commensalism is shown in the relationship of
birds and trees); predation, which is the hunting, killing, and eating of one
species by another (examples: cats and mice; dogs and deer); and competi-
tion, which is defined as an active struggle for survival among all the spe-
cies in a given environment.
This struggle involves the acquisition of various resources: food, terri-
tory, and mates. Food is an obvious target of competition. All organisms
must have energy in order to conduct the cellular chemical reactions (such
as respiration) that keep them alive. Photoautotrophic organisms (plants,
phytoplankton, photobacteria) obtain this energy by converting sunlight,
carbon dioxide, and water into sugar, a process called photosynthesis.
Photoautotrophs, also called producers, compete for light and water. For
example, oak and hickory trees grow taller than most pines, thereby shad-
ing out smaller species and eventually dominating a forest. All other or-
ganisms—animals, zooplankton, and fungi—are heterotrophs; they must
consume other organisms to obtain energy. Heterotrophs include herbi-
vores, carnivores, omnivores, and saprotrophs. Herbivores (plant eaters
such as rabbits and cattle) obtain the sugar manufactured by plants. Carni-
111
Competition
vores (meat eaters such as cats and dogs) eat other heterotrophs in order to
get the sugar that these heterotrophs received from other organisms. Om-
nivores, such as humans, eat plants and animals for the same reason.
Saprotrophs (such as fungi and bacteria) decompose dead organisms for
the same reason. Life on earth functions by intricately complex food chains
in which organisms consume other organisms in order to obtain energy.
Each human being is composed of molecules that were once part of other
living organisms, even other humans. Ultimately, the earth’s energy comes
from the sun.
Territoriality is equally important for two reasons: An organism needs a
place to live, and this place must contain adequate food and water re-
serves. A strong, well-adapted organism will fight and drive away weaker
individuals of the same or different species in order to maintain exclusive
rights to an area containing a large food and water supply. Species that are
less well adapted will be relegated to areas where food and water are
scarce. The stronger species will have more food and will tend to produce
more offspring, since they will easily attract mates. Being stronger or more
adapted does not necessarily mean being physically stronger. A physically
strong organism can be overwhelmed easily by numerous weak individu-
als. In general, adaptability is defined by an organism’s ability to prosper
in a hostile environment and leave many viable offspring.
Animals compete for territory, social status, food, and access to mates. Although
competition may be violent and result in injury, competitive behavior (unlike pre-
dation) rarely results in death. (Corbis)
112
Competition
Types of Competition
Intraspecific competition occurs among individual members of the same
population, for example, when sprouts from plants grow from seeds scat-
tered closely together on the ground. Some seedlings will be able to grow
faster than others and will inhibit the growth of less vigorous seedlings by
overshadowing or overcrowding them.
Within animal species, males attempt to attract females to their territory,
or vice versa, by courtship dances and displays, often including bright col-
ors such as red and blue and exaggerated body size. Mating displays are
very similar to the threat displays used to drive away competitors, al-
though there is no hostility involved. Generally, females are attracted to
dominant males having the best, not necessarily the largest, territories.
Interspecific competition involves two or more different species trying
to use the same resources. All green plants, for example, depend on photo-
synthesis to derive the energy and carbon they need. Different areas or
communities favor different growth characteristics. For plants with high
light requirements, a taller-growing plant (or one with more or broader
leaves) will have a competitive advantage if its leaves receive more direct
sunlight than competitors. If, on the other hand, the species cannot tolerate
much sun, a shorter-growing species that can benefit from sheltering shad-
ows of larger plants nearby will have the competitive advantage over other
shade-loving plants.
Competition for food and territory is both interspecific and intra-
specific. Competition for mates is intraspecific. In an environment, the
place where an organism lives (such as a eucalyptus tree or in rotting logs)
is referred to as its habitat. Simultaneously, each species has its own unique
niche, or occupation, in the environment (such as decomposer or carni-
vore). More than one species can occupy a habitat if they have different
ecological niches. When two or more different species occupy the same
habitat and niche, competition arises. One species will outcompete and
dominate, while the losing competitors may become reduced in numbers
and may be driven away from the habitat.
Pecking Orders
In vertebrate organisms, intraspecific competition occurs between males as
a group and between females as a group. Rarely is there male-versus-female
competition, except in species having high social bonding—primates, for
example. Competition begins when individuals are young. During play
fighting, individuals nip or peck at each other while exhibiting threat dis-
plays. Dominant individuals exert their authority, while weaker individu-
als submit. The net result is a very ordered ranking of individuals from top
113
Competition
to bottom, called a dominance hierarchy or pecking order. The top individ-

ual can threaten and force into submission any individual below it. The
number two individual can threaten anyone except number one, and so
on. The lowest-ranked individual can threaten no one and must submit to
everyone. The lowest individual will have the least food, worst territory,
and fewest (if any) mates. The number one individual will have the most
food, best territory, and most mates. The pecking order changes over time
because of continued group competition that is shown by challenges, ag-
ing, and accidents.
Pecking orders are evident in hens. A very dominant individual will
peck other hens many times but will rarely be pecked. A less dominant in-
dividual will peck less but be pecked more. A correct ranking can be ob-
tained easily by counting the pecking rate for each hen.
In the Netherlands, male black grouse contend with one another in an
area called a “lek,” which may be occupied by as many as twenty males.
The males establish their territories by pecking, wing-beating, and threat
displays. The most dominant males occupy small territories (several hun-
dred meters) at the center of the lek, where the food supply is greatest. Less
dominant males occupy larger territories with less food reserves to the ex-
terior of the lek. Established territories are maintained at measurable dis-
tances by crowing and flutter-jumping, with the home territory owner
nearly always winning. Females, which nest in an adjoining meadow, are
attracted to dominant males in the heavily contested small central territo-
ries.
A baboon troop can range in size from ten to two hundred members, but
usually averages about forty. Larger, dominant males and their many fe-
male mates move centrally within the troop. Less dominant males, with
fewer females, lie toward the outside of the troop. Weak individuals at the
troop periphery are more susceptible to predator attacks. Dominant males
exert their authority by threat displays, such as the baring of the teeth or
charging; weaker males submit by presenting their hindquarters. Conflicts
are usually peacefully resolved.
Female lions maintain an organized pride with a single ruling male.
Young males are expelled and wander alone in the wilderness. Upon
reaching adulthood, males attempt to take over a pride in order to gain ac-
cess to females. If a male is successful in capturing a pride and expelling his
rival, he will often kill the cubs of the pride, simultaneously eliminating his
rival’s descendants and stimulating the females to enter estrus for mating.
Competition Within Niches

Interspecific competition occurs between different species over food and
114
Competition
water reserves and territories. Two or more species occupying the same
niche and habitat will struggle for the available resources until either one
species dominates and the others are excluded from the habitat or the dif-
ferent species evolve into separate niches by targeting different food re-
serves, thus enabling all to survive in the same habitat. Numerous inter-
specific studies have been conducted—on crossbills, warblers, blackbirds,
and insects, to mention a few.
Crossbills are small birds that live in Europe and Asia. Three crossbill
species inhabit similar habitats and nearly similar niches. Each species has
evolved a slightly modified beak, however, for retrieving and eating seeds
from three different cone-bearing (coniferous) trees. The white-winged
crossbill has a slender beak for feeding from small larch cones, the com-
mon crossbill has a thicker beak for feeding from larger spruce cones, and
the parrot crossbill appropriately has a very thick beak for feeding from
pine cones. The evolution of different niches has enabled these three com-
petitors to survive.
Another example of this phenomenon is shown by five species of
warblers that inhabit the coniferous forests of the American northeast.
The myrtle warbler eats insects from all parts of trees up to seven meters
high. The bay-breasted warbler eats insects from tree trunks six to twelve
meters above the ground. The black-throated green, blackburnian, and
cape may warblers all feed near the treetops, according to elaborate stud-
ies by Robert H. MacArthur. The coexistence of five different species
is probably the result of the warblers occupying different parts of the
trees, with some warblers developing different feeding habits so that all
survive.
G. H. Orians and G. Collier studied competitive exclusion between red-
wing and tricolored blackbirds. Introduction of tricolored blackbirds into
redwing territories results in heavy redwing aggression, although the
tricolored blackbirds nearly always prevail.
Two species of African ants, Anoplolepis longipes and Oecophylla longi-
noda, fight aggressively for territorial space. M. J. Way found that Anoplolepis
prevails in sandy environments, whereas Oecophylla dominates in areas
having thick vegetation.
Interspecific competition therefore results in the evolution of new traits
and niches and the exclusion of certain species. Mathematical models of
competition are based upon the work of A. J. Lotka and V. Volterra. The
Lotka-Volterra equations attempt to measure competition between species
for food and territory based upon the population size of each species, the
density of each species within the defined area, the rate of population in-
crease of each species, and time.
115
Competition
Observing Competiton
Studies of competition between individuals of the same or different spe-
cies generally follow one basic method: observation. Interactions between
organisms are observed and carefully measured to determine if the situa-
tion is competition, predation, parasitism, or mutualism. More detailed
analyses of environmental chemical and physical conditions are used
to determine the existence of additional influences. Observations of com-
petition between organisms involve direct visual contact in the wild,
mark-recapture experiments, transplant experiments, measurements of
population sizes in given areas, and competition experiments in artificial
environments.
Direct visual contact involves the scientist entering the field, finding a
neutral, nonthreatening position, and watching and recording the actions
of the subject organisms. The observer must be familiar with the habits of
the subject organism and must be keen to detect subtle cues such as facial
gestures, vocalizations, colors, and patterns of movement from individual
to individual. Useful instruments include binoculars, telescopes, cameras,
and sound recorders. The observer must be capable of tracking individuals
over long distances so that territorial boundaries and all relevant actions
are recorded. The observer may have to endure long periods of time in the
field under uncomfortable conditions.
Mark-recapture experiments involve the capture of many organisms,
tagging them, releasing them into an area, and then recapturing them
(both tagged and untagged) at a later time. Repeated collections (recap-
tures) over time can give the experimenter an estimate of how well the spe-
cies is faring in a particular environment. This technique is used in con-
junction with other experiments, including transplants and population
size measurements.
In transplant experiments, individuals of a given species are marked
and released into a specific environmental situation, such as a new habitat
or another species’ territory. The objective of the experiment is to see
how well the introduced species fares in the new situation, as well as the
responses of the various species which normally inhabit the area. The
tricolored blackbird takeover of redwing blackbird territories is a prime
example. Another example is the red wolf, a species that was extinct in the
wild until several dozen captive wolves were released at the Alligator
River Wildlife Refuge in eastern North Carolina. Their survival is uncer-
tain. Accidental transplants have had disastrous results for certain species;
for example, the African honeybee poses a threat to the honey industry in
Latin America and the southern United States because it is aggressive and
produces poorly.
116
Competition
Measurements of population sizes rely upon the point-quarter tech-

nique, in which numerous rectangular areas of equal size are marked in the
field. The number of organisms of each species in the habitat is counted for
a given area; an averaging of all areas is then made to obtain a relatively
accurate measure of each population’s size. In combination with mark-
recapture experiments, population measurements can provide informa-
tion for birthrates, death rates, immigration, and emigration over time for
a given habitat.
Laboratory experiments involve confrontations between different spe-
cies or individuals of the same species within an artificial environment. For
example, male mouse (Mus musculus) territoriality can be studied by intro-
ducing an intruder into another male’s home territory. Generally, the win-
ner of the confrontation is the individual that nips its opponent more
times. Usually, home court advantage prevails; the intruder is driven
away. Similar studies have been performed with other mammalian, reptile,
fish, insect, and bird species.
Interactions between different species are subtle and intricate. Seeing
how organisms associate enables scientists to understand evolution and to
model various environments. Competition is a major driving force in evo-
lution. The stronger species outcompete weaker species for the available
ecological niches. Mutations in organisms create new traits and, therefore,
new organisms (more species), which are selected by the environment for
adaptability.
All environments consist of a complex array of species, each dependent
on the others for survival. The area in which they live is their habitat. Each
species’ contribution to the habitat is that species’ niche. More than one
species in a given habitat causes competition. Two species will struggle for
available territory and food resources until either one species drives the
other away or they adapt to each other and evolve different feeding habits
and living arrangements. Competition can be interspecific (between indi-
viduals of different species) or intraspecific (between individuals of the
same species). The environment benefits because the most adapted species
survive, whereas weaker species are excluded.
David Wason Hollar, Jr.
See also: Allelopathy; Animal-plant interactions; Biodiversity; Biogeog-

raphy; Biological invasions; Coevolution; Communities: ecosystem inter-
actions; Communities: structure; Defense mechanisms; Food chains and
webs; Gene flow; Genetic diversity; Lichens; Mycorrhizae; Pollination;
Predation; Speciation; Succession; Symbiosis; Trophic levels and ecological
niches.
117
Competition

Andrewartha, H. G. Introduction to the Study of Animal Populations. Chicago:
University of Chicago Press, 1967.
Arthur, Wallace. The Niche in Competition and Evolution. New York: John
Wiley & Sons, 1987.
Hartl, Daniel L. Principles of Population Genetics. 3d ed. Sunderland, Mass.:
Sinauer Associates, 1997.
Keddy, Paul A. Competition. 2d ed. New York: Kluwer, 2000.
Lorenz, Konrad. On Aggression. New York: Harcourt, Brace & World, 1963.
Raven, Peter H., and George B. Johnson. Biology. 5th ed. Boston: WCB/
McGraw-Hill, 1999.
Wilson, Edward O. Sociobiology: The New Synthesis. Cambridge, Mass.:
Belknap Press of Harvard University Press, 1975.
118
CONSERVATION BIOLOGY
Type of ecology: Restoration and conservation ecology
Conservation biology is a multidisciplinary field that uses knowledge and skills

from all aspects of biological science to design and implement methods to ensure
the survival of species, ecosystems, and ecological processes.
T he overriding mission of conservation biology is to ensure the contin-

ued survival of all life-forms and to maintain the structure and func-
tion of all ecosystems and ecological processes. Attempting to achieve this
goal requires biological knowledge from disciplines such as genetics,
physiology, systematics, and ecology in combination with skills and prac-
tices from applied fields such as forestry, fisheries, and wildlife manage-
ment. In addition to biological knowledge, social and economic issues
must be addressed in the development of natural resource policies de-
signed to preserve and protect species and ecosystems.
Conservation biology, a relatively new discipline, arose from questions
about how to develop methods and policies to maintain global biodiver-
sity. Policy development and implementation must consider the role of
evolutionary processes in order to preserve genetic diversity and enable
the continued survival of threatened or endangered species. Conservation
biologists are also concerned with practical issues such as the design of
parks and nature reserves.
David D. Reed
See also: Biodiversity; Deforestation; Endangered animal species; Endan-

gered plant species; Erosion and erosion control; Forest management; Ge-
netic diversity; Grazing and overgrazing; Integrated pest management;
Landscape ecology; Multiple-use approach; Old-growth forests; Refores-
tation; Restoration ecology; Species loss; Sustainable development; Urban
and suburban wildlife; Waste management; Wildlife management; Zoos.

Cox, G. W. Conservation Biology: Concepts and Applications. Dubuque, Iowa:
William C. Brown, 1997.
Hunter, Malcolm, Jr. Fundamentals of Conservation Biology. Cambridge,
Mass.: Blackwell Science, 1996.
Primack, Richard. Essentials of Conservation Biology. Sunderland, Mass.:
119
CONVERGENCE AND DIVERGENCE
Some of the most dramatic examples of natural selection are the result of adapta-
tion in response to stressful climatic conditions. Such selection may cause unre-
lated species to resemble one another in appearance and function, a phenomenon
known as convergence. In other situations, subpopulations of a single species may
split into separate species as the result of natural selection. Such divergence is best
seen on isolated islands.
Convergent Evolution
Convergent evolution occurs when organisms from different evolutionary
lineages evolve similar adaptations to similar environmental conditions.
This can happen even when the organisms are widely separated geograph-
ically. A classic example of convergent evolution occurred with Cactaceae,
the cactus family, of the Americas and with the spurge or euphorbs
(Euphorbiaceae) family of South Africa, both of which have evolved succu-
lent (water-storing) stems in response to desert conditions.
The most primitive cacti are vinelike, tropical plants of the genus
Pereskia. These cacti, which grow on the islands of the West Indies and in
tropical Central and South America, have somewhat woody stems and
broad, flat leaves. As deserts developed in North and South America,
members of the cactus family began to undergo selection for features that
were adaptive to hotter, dryer conditions.
The stems became greatly enlarged and succulent as extensive water-
storage tissues formed in the pith or cortex. The leaves became much re-
duced. In some cactus species, such as the common prickly pear (Opuntia),
the leaves are small, cylindrical pegs that shrivel and fall off after a month
or so of growth. In most cacti, only the leaf base forms and remains as a
small hump of tissue associated with an axillary bud. In some cacti this
hump is enlarged and is known as a tubercle. Axillary buds in cacti are
highly specialized and are known as areoles. The “leaves” of an areole are
reduced to one or more spines. Particularly in columnar cacti, the areoles
are arranged in longitudinal rows along a multiple-ridged stem.
With the possible exception of the genus Rhipsalis, which has one spe-
cies reported to occur naturally in Africa, all cacti are native to the Ameri-
cas. As deserts formed in Africa, Eurasia, and Australia, different plant
families evolved adaptations similar to those in cacti. The most notable ex-
amples are the candelabra euphorbs of South Africa. Desert-dwelling
120
Convergence and divergence
members of the Euphorbiaceae frequently have succulent, ridged, cylindri-

cal stems resembling those of cacti. The leaves are typically reduced in size
and are present only during the rainy season. They are arranged in rows
along each of several ridges of the stem. Associated with each leaf are one
or two spines. As a result, when the leaves shrivel and fall off during the
dry season, a spiny, cactuslike stem remains.
The succulent euphorbs of Africa take on all the forms characteristic
of American cacti, from pincushions and barrels to branched and un-
branched columns. Other plant families that show convergence with the
cacti, in having succulent stems or leaves, are the stem succulents of the
milkweed family, sunflower family, stonecrop family, purslane family,
grape family, leaf succulents of the ice plant family, daffodil family, pineap-
ple family, geranium family, and lily family.
Divergent Evolution
Some of the most famous examples of divergent evolution have occurred
in the Galápagos Islands. The Galápagos comprise fourteen volcanic is-
lands located about 600 miles west of South America. A total of 543 species
of vascular plants are found on the islands, 231 of which are endemic,
found nowhere else on earth. Seeds of various species arrived on the is-
lands by floating in the air or on the water or being carried by birds or hu-
mans.
With few competitors and many different open habitats, variant forms
of each species could adapt to specific conditions, a process known as
An example of convergent evolution is the way that sharks (left), which are fish,
and dolphins (right), which are mammals, have evolved similar body shapes to
adapt to their marine ecological niches. Although the two animals look very much
alike, their differences in evolutionary terms are vast. (Digital Stock)
121
Convergence and divergence
adaptive radiation. Those forms of a species best suited to each particular

habitat were continually selected for and produced progeny in that habi-
tat. Over time, this natural selection resulted in multiple new species shar-
ing the same ancestor. The best examples of divergent evolution in the
Galápagos have occurred in the Cactaceae and Euphorbiaceae. Eighteen spe-
cies and variety of cacti are found on the islands, and all are endemic. Of
the twenty-seven species and varieties of euphorbs, twenty are endemic.
An interesting example of the outcome of divergent evolution can be
seen in the artificial selection of different cultivars (cultivated varieties) in
the genus Brassica. The scrubby Eurasian weed colewort (Brassica oleracea)
is the ancestor of broccoli, brussels sprouts, cabbage, cauliflower, kale, and
kohlrabi (rutabaga). All these vegetables are considered to belong to the
same species, but since the origin of agriculture, each has been selected for
a specific form that is now recognized as a distinct crop.
See also: Adaptations and their mechanisms; Adaptive radiation; Coevo-

lution; Colonization of the land; Dendrochronology; Development and
ecological strategies; Evolution: definition and theories; Evolution: his-
tory; Evolution of plants and climates; Extinctions and evolutionary explo-
sions; Gene flow; Genetic drift; Genetically modified foods; Isolating
mechanisms; Natural selection; Nonrandom mating, genetic drift, and
mutation; Paleoecology; Population genetics; Punctuated equilibrium vs.
gradualism; Speciation; Species loss.

Bowman, Robert I., Margaret Berson, and Alan E. Leviton. Patterns of Evo-
lution in Galápagos Organisms. San Francisco: California Academy of Sci-
ences, 1983.
Darwin, Charles. “Journal and Remarks: 1832-1836.” In Narrative of the Sur-
veying Voyages of His Majesty’s Ships Adventure and Beagle Between the
Years 1826 and 1836: Describing Their Examination of the Southern Shores of
South America, and the Beagle’s Circumnavigation of the Globe, edited by
Robert Fitzroy. Vol. 3. Reprint. New York: AMS Press, 1966.
Harris, James G., and Melinda Woolf Harris. Plant Identification Terminol-
ogy: An Illustrated Glossary. Spring Lake, Utah: Spring Lake, 1994.
Uno, Gordon, Richard Storey, and Randy Moore. Principles of Botany. New
York: McGraw-Hill, 2001.
122
DEEP ECOLOGY
Type of ecology: Theoretical ecology
Deep ecology is a school of environmental philosophy based on environmental ac-

tivism and ecological spirituality.
T he term “deep ecology” was first used by Norwegian philosopher

Arne Naess in 1972 to suggest the need to go beyond the anthropocen-
tric view that nature is merely a resource for human use. The concept has
been used in three major ways. First, it refers to a commitment to deep
questioning about environmental ethics and the causes of environmental
problems. Such questioning leads to critical reflection on the fundamental
worldviews that underlie specific environmental ideas and practices. Sec-
ond, deep ecology refers to a platform of generally agreed upon values that
a variety of environmental activists share. These values include an affirma-
tion of the intrinsic value of nature, the recognition of the importance of
biodiversity, a call for a reduction of human impact on the natural world,
greater concern with quality of life rather than material affluence, and a
commitment to changing economic policies and the dominant view of na-
ture. Third, deep ecology refers to particular philosophies of nature that
tend to emphasize the value of nature as a whole (ecocentrism), an identifi-
cation of the self with the natural world, and an intuitive and sensuous
communion with the earth.
Because of its emphasis on fundamental worldviews, deep ecology is
often associated with non-Western spiritual traditions such as Buddhism
and Native American cultures, as well as radical Western philosophers such
as Baruch Spinoza and Martin Heidegger. It has also drawn on the nature
writing of Henry David Thoreau, John Muir, Robinson Jeffers, and Gary
Snyder. Deep ecology’s holistic tendencies have led to associations with
the Gaia hypothesis, and its emphasis on diversity and intimacy with nature
has linked it to bioregionalism. Deep ecological views have also had a strong
impact on environmental activism, including the Earth First! movement.
Deep ecologists have sometimes criticized the animal rights perspective
for continuing the traditional Western emphasis on individuals while ne-
glecting whole systems, as well as for a revised speciesism that still values
certain parts of nature (animals) over others. Some deep ecologists have
also been critical of mainstream environmental organizations such as the
Sierra Club for not confronting the root causes of environmental degrada-
tion.
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Deep ecology
On the other hand, deep ecology has been criticized by ecofeminists for
failing to consider gender differences in the experience of the self and na-
ture, the lack of an analysis of the tie between the oppression of women
and nature, and promoting a holism that supposedly disregards the reality
and value of individuals and their relationships. Social ecologists have crit-
icized deep ecology for a failure to critique the relationship between envi-
ronmental destruction on the one hand and social structure and political
ideology on the other. In addition, a distrust of human interference with
nature has led some thinkers to present the ideal as pristine wilderness
with no human presence. In rare and extreme cases, deep ecologists have
implied a misanthropic attitude. In some instances, especially early writ-
ings by deep ecologists, such criticisms have considerable force. However,
these problematic views are not essential to deep ecology, and a number of
thinkers have developed a broadened view that overlaps with ecofem-
inism and social ecology.
David Landis Barnhill
See also: Balance of nature; Biomes: determinants; Biomes: types; Bio-

sphere concept; Ecology: definition; Ecosystems: definition and history;
Sustainable development; .

Naess, Arne. Ecology, Community, and Lifestyle: Outline of an Ecosophy.
Translated and revised by David Rothenberg. New York: Cambridge
_______. Spinoza and the Deep Ecology Movement. Delft: Eburon, 1993.
124
DEFENSE MECHANISMS
Types of ecology: Behavioral ecology; Chemical ecology; Physiological
ecology
All organisms represent a potential resource for their predators. Several have
evolved ingenious ways to prevent themselves from becoming a predator’s next
meal.
A ll organisms are composed of fixed carbon, biomolecules, and min-

eral nutrients and therefore represent energy and nutrient resources
for consumers. To be successful in life, animals must avoid, tolerate, or de-
fend themselves against natural enemies such as predators, parasites, and
competitors. The term “defense” can be attributed to any trait that reduces
the likelihood that an organism, or part of an organism, will be consumed
by a predator. Several categories of defenses have evolved in animals, in-
cluding structural defenses, chemical defenses, associational defenses, be-
havioral defenses, autotomy, and nutritional defenses. Animals often pos-
sess more than one type of defense, thereby having backup plans in case
the first line of defense fails. The number of defenses devised by organisms
is a reflection of the strong selective pressure exerted by predators.
Structural Defenses: Being Hard and Sharp

Structures that defend animals can act as external shields: sharp spines
located externally or internally, skeletal materials that make tissues too
hard to bite easily, or weaponry such as horns, teeth, and claws. External
structures that protect vulnerable soft tissues include the chitonous exo-
skeletons of crustaceans, the calcareous shells of corals, mollusks, and bar-
nacles, the tests (skeletal plates) of echinoderms, the tough tunics of ascid-
ians, and the hard plates of armadillos. The pretty shells that tourists
collect along beaches were once used to protect a soft, delicate animal that
lived inside the shell. Hard, protective shells remain after the animal dies
and can be used by other animals for protection. For example, small fish
will retreat into empty conch shells when they feel threatened by preda-
tors, and hermit crabs live inside empty snail shells to protect their soft,
vulnerable abdomens.
Some animals cover their bodies with sharp structures that puncture
predators that try to bite them. The porcupine is a good example of a mam-
mal that uses this defensive strategy. Porcupines are covered with tens of
thousands of long, pointed spines, or quills, growing from their back and
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Defense mechanisms
sides. The quills have needle-sharp ends containing hundreds of barbs that
make the quills difficult to remove. Sea urchins are also covered with long,
sharp spines that deter would-be predators. Urchins can move their spines,
and will direct them toward anything that comes in contact with them,
such as a predator. While porcupines and urchins are covered with multi-
ple spines, stingrays defend themselves from enemies by inflicting a wound
with a single barbed spine. The wound is extremely painful, giving these
rays their common name.
Predators have sharp claws and teeth that help them grasp, subdue, and
consume their prey. These same structures, used offensively in hunting,
can also be used to protect themselves from their own predators. Small
predators such as badgers, raccoons, and foxes can fend off larger preda-
tors such as wolves and mountain lions with their weaponry. Rather than
risk injury, the larger predators will avoid a fight with the smaller predator
and seek a less risky meal, such as a rabbit or mouse.
Chemical Defenses: Poor Taste, Bad Smell, or Toxic Chemicals

Both plants and animals defend themselves by using compounds that are
distasteful, toxic, or otherwise repulsive to consumers. Most defensive
compounds are secondary metabolites of unique structures, but can also
include more generic compounds such as sulfuric acid or calcium carbon-
ate. Secondary metabolites get their name because they are not involved
in basic metabolic pathways such as respiration or photosynthesis (that
is, primary metabolic reactions), not because they are of secondary im-
portance. Indeed, many organisms probably could not survive in their
natural environment without the protection of their secondary metabo-
lites.
Stink bugs get their names because of the smelly secondary metabolites
they release from pores located on the sides of their thorax. These smelly
compounds repel predators, and may even indicate toxicity to the preda-
tor. These insects are common garden pests that are usually controlled
with chemical pesticides. However, it appears that the eggs of stink bugs
are not defended against roly-poly pill bugs, which can control stink bug
numbers (and hence, garden damage) by preying on eggs.
Bombardier beetles take chemical defenses a step further, erupting a
boiling hot spray of chemicals in the direction of a predator. To accomplish
this, the bombardier beetle has a pair of glands that open at the tip of its ab-
domen. Each gland has two compartments, one that contains a solution of
hydroquinone and hydrogen peroxide, and the other that contains a mix-
ture of enzymes. When threatened by a predator, the bombardier beetle
squeezes the hydroquinone and hydrogen peroxide mixture into the en-
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Defense mechanisms
zyme compartment, where an exothermic reaction that produces quinone

takes place. The large amount of heat generated brings the quinone mix-
ture to its boiling point, and it is forcefully emitted as a vapor toward the
threat. An average bombardier beetle can produce about twenty loud dis-
charges of repulsive, hot chemicals in quick succession.
Chemical defenses are common among small, slow animals such as in-
sects, sponges, cnidarians, and sea slugs, which might be limited in their
ability to flee from predators. However, chemical defenses are rather rare
among large, fast animals. One of the few mammals that uses chemical de-
fenses is the black-and-white-striped skunk. Most people are familiar with
the smelly chemical brew emitted from these animals, as it is distinctly de-
tectable along roads when skunks get hit by cars, and can be detected up to
a mile from the location where a skunk sprays. These mammals hold their
smelly musk in glands located below their tail, and squirt the liquid
through ducts that protrude from the anus. When threatened by a preda-
tor, the skunk raises its tail and directs its rear end toward the predator. A
predator that has had prior experience with a skunk might retreat from this
display, but if the predator is persistent at harassing the skunk, the striped
mammal will deliver a spray of smelly chemicals that usually sends the
predator running. The musk also causes intense pain and temporary blind-
ness if it gets in the eyes of the predator.
Associational Defenses
Associational defenses occur when a species gains protection from a natu-
ral enemy by associating with a protective species, such as when humans
gain protection from enemies by keeping a guard dog on their property.
Types of protection provided to the defended species through this coevolu-
tion can be structural, chemical, or aggressive.
Small animals can avoid predators by using a defended species as habi-
tat. For example, small fish defend themselves by associating with sea ur-
chins, gaining protection by hiding among the sharp spines. Some species
of shrimp inhabit the cavities and canals of sponges. Sponges are known to
be chemically and structurally defended against most predators, with the
exception of angel fish and parrot fish. Finally, much of the diverse coral
reef fauna seek protection among the cracks and the crevices in the reef.
Reefs, slowly built by coral animals, are the largest structures ever made by
living organisms, and serve a protective role for thousands of species that
inhabit reefs.
Associational defenses can also be chemically mediated. For example,
bacteria that grow symbiotically on shrimp eggs produce secondary me-
tabolites that protect the egg from a parasitic fungus. The numerous ex-
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Defense mechanisms
amples of sequestration of chemical defenses can be categorized as asso-

ciational defenses, as they involve associating with chemically defended
prey.
An organism might even be defended by protective species that aggres-
sively attack would-be predators, especially if the protected species is a re-
source for the aggressive defender. For example, humans are protected by
guard dogs because dogs view people as a resource that provides them
with food, water, and shelter. Stop feeding the dog, and it is likely to look
elsewhere for somebody to protect. There are several nonhuman examples
of aggressive defensive associations, especially among ants. Aphids are in-
sects that feed on the sugary phloem stream of plants. In the process of
feeding and processing phloem, the aphids secrete large amounts of hon-
eydew, which the ants harvest and consume; that is, aphids provide ants
with a resource. Ants tend to aphids in the same way that dairy farmers
tend to their cows. The ants carry aphids to prime feeding locations, de-
fend aphids from predators, and periodically “milk” the aphids of their
honeydew by stroking them with their antennae.
Aposematic Coloration and Mimicry

Being chemically defended does not protect an animal from being acciden-
tally eaten. Therefore, chemically defended animals often advertise the
fact that they are nasty to avoid such accidents. This advertisement is often
in the form of outlandish colors and patterns that flaunt the animal’s dis-
tastefulness to predators. Using bright warning patterns is called apose-
matic coloration.
One problem with aposematic coloration is the training of predators:
Bright coloration is useful only if the predator understands the warning.
Otherwise, the coloration simply makes the animal a conspicuous prey
item. One way that different species with aposematic coloration share the
cost of training naïve predators is through mimicry. A predator that eats an
individual of species A (assume species A is bright red with blue stripes)
and vomits shortly thereafter may learn to avoid things that are red with
blue stripes, though at the cost of that first individual’s life. This educated
predator will now avoid other members of species A, and any other organ-
ism that looks like species A (the mimic), whether the mimic is toxic or not.
If the mimic is toxic, the system is termed Müllerian mimicry. If the mimic
is a palatable species that looks like a toxic model, the system is termed
Batesian mimicry.
Mimicry is common within groups of closely related organisms (for ex-
ample, snakes, butterflies, and bees) which are already similar in appear-
ance. However, mimicry can also occur even when the model and mimic
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Defense mechanisms
are distantly related. For example, there are caterpillars that mimic the
head of a snake, moths that mimic the eyes of a cat, and beetles, moths, and
flies that mimic stinging bees and wasps.
Autotomy: Throw the Predator a Bone

Sometimes, despite the best defenses, a predator will get hold of a prey.
When this happens, some animals are able to sacrifice a portion of their
body to the predator, with the hopes that the remaining parts will survive,
and perhaps even regrow the lost parts. This ability to lose a body part in-
tentionally is called autotomy.
Many lower animals, such as sponges, cnidarians, and worms, have
great regeneration abilities, and can regrow body parts well. In fact, these
animals can even use regeneration as a form of asexual reproduction:
Break the animal into four parts, and the parts will generate four complete
individuals.
Sea cucumbers, in addition to being chemically defended, are able to
eviscerate (autotomy of intestines) when harassed by a predator. These are
not fast animals, so this action does not allow them to escape, but it might
satisfy (or disgust) the predator enough to make it lose interest in the sea
cucumber. Losing a large portion of its digestive tract interferes with feed-
ing, but the sea cucumber can regenerate those parts of the gut that were
eviscerated, restoring itself to original function. Sea cucumbers also play
an important role in a defensive association with the pearlfish. When the
pearlfish feels threatened, it locates the anus of a sea cucumber, then backs
into its intestine, where it hides until the danger has passed.
The regenerative ability of higher animals is generally less than that of
lower animals. However, autonomy does occur even in some vertebrates.
Lizards are well known for their ability to release the tips of their tails
when grabbed by a predator. The predator is distracted, and perhaps satis-
fied, by the wiggling piece of flesh, and in the meantime, the remainder of
the lizard scampers off to safety. Geckos release skin instead of tails. The
part of the skin that is grabbed by the predator is released, enabling the
gecko to break free and escape.
Nutritional Defenses
Some animals, such as corals, jellyfish, anemones, and gorgonians (phy-
lum Cnidaria), possess a type of combined structural and chemical de-
fense in the form of specialized stinging cells called nematocysts. When
nematocysts are stimulated, they rapidly discharge a barb that punctures
the skin of a predator, often releasing toxic chemicals at the same time. The
stinging sensation that people get when they come into contact with a jelly-
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Defense mechanisms
fish is caused by nematocysts. Some of these jellyfish stings are so potent

that they can result in death.
Not only do many predators avoid jellyfish because they posses nem-
atocysts, but predators may avoid jellyfish because they are jellylike, being
composed of more than 95 percent water. It takes time and effort for preda-
tors to locate, handle, ingest, and digest prey. If the prey item is basically a
bag of seawater (as jellyfish are), then predators might not bother eating
these nutrient-deficient animals. Thus, these animals are “nutritionally”
defended. Nutritional defenses are also used by plants, but they are gener-
ally not an available strategy for animals other than jellyfish, as most ani-
mal tissue is relatively nutritious.
Greg Cronin
See also: Allelopathy; Animal-plant interactions; Bioluminescence; Co-

evolution; Communities: ecosystem interactions; Metabolites; Poisonous
animals; Poisonous plants; Predation; Territoriality and aggression.

Cloudsley-Thompson, John L. Tooth and Claw: Defensive Strategies in the An-
imal World. London: J. M. Dent & Sons, 1980.
Edmunds, Malcolm. Defence in Animals. Burnt Mill, England: Longman,
1974.
Evans, David L., and Justin O. Schmidt, eds. Insect Defenses: Adaptive Mech-
anisms and Strategies of Prey and Predators. Albany: State University of
New York Press, 1990.
Kaner, Etta. Animal Defenses: How Animals Protect Themselves. Toronto: Kids
Can Press, 1999.
McClintock, James B., and Bill J. Baker, eds. Marine Chemical Ecology. Boca
Raton, Fla.: CRC Press, 2001.
Owen, Denis. Survival in the Wild: Camouflage and Mimicry. Chicago: Uni-
versity of Chicago Press, 1980.
130
DEFORESTATION
Types of ecology: Ecotoxicology; Restoration and conservation ecology
Deforestation is the loss of forestlands through encroachment by agriculture, in-

dustrial development, or nonsustainable commercial forestry, and other human as
well as natural activity.
C oncerns about deforestation, particularly in tropical regions, have

risen as the role that tropical rain forests play in moderating global cli-
mate has become better understood. Environmental activists decried the
apparent accelerating pace of deforestation in the twentieth century be-
cause of the potential loss of wildlife and plant habitat and the negative ef-
fects on biodiversity. By the 1990’s research by mainstream scientists had
confirmed that deforestation was indeed occurring on a global scale and
that it posed a serious threat to global ecology.
Deforestation as a result of expansion of agricultural lands or nonsus-
tainable timber harvesting has occurred in many regions of the world at
different periods in history. The Bible, for example, refers to the cedars of
Lebanon. Lebanon, like many of the countries bordering the Mediterra-
nean Sea, was thickly forested several thousand years ago. A growing hu-
man population, overharvesting, and the introduction of grazing animals
such as sheep and goats decimated the forests, which never recovered.
Countries in Latin America, Asia, and Africa have also lost woodlands.
While some of this deforestation is caused by a demand for tropical hard-
woods for lumber or pulp, the leading cause of deforestation in the twen-
tieth century, as it was several hundred years ago, was the expansion of
agriculture. The growing demand by the industrialized world for agricul-
tural products such as beef has led to millions of acres of forestland being
bulldozed or burned to create pastures for cattle. Researchers in Central
America have watched with dismay as large beef-raising operations have
expanded into fragile ecosystems in countries such as Costa Rica, Guate-
mala, and Mexico.
A tragic irony in this expansion of agriculture into tropical rain forests is
that the soil underlying the trees is often unsuited for pastureland or rais-
ing other crops. Exposed to sunlight, the soil is quickly depleted of nutri-
ents and often hardens. The once-verdant land becomes an arid desert,
prone to erosion, that may never return to forest. As the soil becomes less
fertile, hardy weeds begin to choke out the desirable forage plants, and the
cattle ranchers move on to clear a fresh tract.
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Deforestation
Slash-and-Burn Agriculture
Beef industry representatives often argue that their ranching practices are
simply a form of slash-and-burn agriculture and do no permanent harm. It
is true that many indigenous peoples in tropical regions have practiced
slash-and-burn agriculture for millennia, with only a minimal impact on
the environment. These farmers burn shrubs and trees to clear small plots
of land.
Anthropological studies have shown that the small plots these peasant
farmers clear can usually be measured in square feet, not hectares as cattle
ranches are, and are used for five to ten years. As fertility declines, the
farmer clears a plot next to the depleted one. The farmer’s family or village
will gradually rotate through the forest, clearing small plots and using
them for a few years, and then shifting to new ground, until they eventu-
ally come back to where their ancestors began one hundred or more years
before.
As long as the size of the plots cleared by farmers remains small in pro-
portion to the forest overall, slash-and-burn agriculture does not contrib-
ute significantly to deforestation. If the population of farmers grows, how-
ever, more land must be cleared with each succeeding generation. In many
tropical countries, traditional slash-and-burn agriculture can then be as
ecologically devastating as the more mechanized cattle ranching opera-
tions.
Logging
Although logging is not the leading cause of deforestation, it is a signifi-
cant factor. Tropical forests are rarely clear-cut by loggers, as they typically
contain hundreds of different species of trees, many of which have no com-
mercial value. Loggers may select trees for harvesting from each stand. Se-
lective harvesting is a standard practice in sustainable forestry. However,
just as loggers engaged in the disreputable practice of high-grading across
North America in the nineteenth century, so are loggers high-grading in
the early twenty-first century in Malaysia, Indonesia, and other nations
with tropical forests.
High-grading is a practice in which loggers cut over a tract to remove
the most valuable timber while ignoring the damage being done to the re-
sidual stand. The assumption is that, having logged over the tract once, the
timber company will not be coming back. This practice stopped in North
America, not because the timber companies voluntarily recognized the
ecological damage they were doing but because they ran out of easily ac-
cessible, old-growth timber to cut. Fear of a timber famine caused logging
companies to begin forest plantations and to practice sustainable forestry.
132
Deforestation
Percentage of Annual Deforestation by Country, 1990-1995
1 percent From 0 to From –1 to Less than No data

or more 1 percent 0 percent –1 percent
Source: United Nations Food and Agriculture Organization
While global satellite photos indicate that significant deforestation has

occurred in tropical areas, enough easily harvested old-growth forest re-
mains in some areas that there is no economic incentive for timber compa-
nies to switch to sustainable forestry.
Logging may also contribute to deforestation by making it easier for ag-
riculture to encroach on forestlands. The logging company builds roads for
use while harvesting trees. Those roads are then used by farmers and
ranchers to move into the logged tracts, where they clear whatever trees
the loggers have left.
Environmental Impacts
Despite clear evidence that deforestation is accelerating, the extent of the
problem remains debatable. The United Nations Food and Agriculture
Organization (FAO), which monitors deforestation worldwide, bases its
statistics on measurements taken from satellite images. These data indi-
cate that between 1980 and 1990, at least 159 million hectares (393 million
acres) of land became deforested. The data also reveal that, in contrast
to the intense focus on Latin America by both activists and scientists,
the most dramatic loss of forestlands occurred in Asia. The defores-
tation rate in Latin America was 7.45 percent, while in Asia 11.42 per-
cent of the forests vanished. Environmental activists are particularly con-
133
Deforestation
cerned about forest losses in Indonesia and Malaysia, two countries where
timber companies have been accused of abusing or exploiting native peo-
ples in addition to engaging in environmentally damaging harvesting
methods.
Researchers outside the United Nations have challenged the FAO’s
data. Some scientists claim the numbers are much too high, while others
provide convincing evidence that the FAO numbers are too low. Few re-
searchers, however, have tried to claim that deforestation on a global scale
is not happening. In the 1990’s the reforestation of the Northern Hemi-
sphere, while providing an encouraging example that it is possible to re-
verse deforestation, was not enough to offset the depletion of forestland in
tropical areas. The debate among forestry experts centers on whether de-
forestation has slowed, and, if so, by how much.
Deforestation affects the environment in a multitude of ways. The most
obvious effect is a loss of biodiversity. When an ecosystem is radically al-
tered through deforestation, the trees are not the only thing to disappear.
Wildlife species decrease in number and in variety. As forest habitat
Results of Deforestation
Population increase
(human and animal)
Requiring more:
Agricultural land Rangeland Forest products
Contributing to:
Deforestation
Resulting in:
Increased soil Loss of Reduced water Reduction in Fuelwood

erosion by shelterbelt retention in biological scarcity
wind and water soil diversity
Increased Desertification Increased Loss of Higher

reservoir flooding endangered socioeconomic
siltation species costs
Source: Adapted from A. K. Biswas, “Envioronmental Concerns in Pakistan, with Special
Reference to Water and Forests,” in Environmental Conservation, 1987.
134
Deforestation
shrinks through deforestation, many plants and animals become vulnera-

ble to extinction. Many biologists believe that numerous animals and
plants native to tropical forests will become extinct from deforestation be-
fore humans have a chance to even catalog their existence.
Other effects of deforestation may be less obvious. Deforestation can
lead to increased flooding during rainy seasons. Rainwater that once
would have been slowed or absorbed by trees instead runs off denuded
hillsides, pushing rivers over their banks and causing devastating floods
downstream. The role of forests in regulating water has long been recog-
nized by engineers and foresters. Flood control was, in fact, one of the mo-
tivations behind the creation of the federal forest reserves in the United
States during the nineteenth century. More recently, disastrous floods in
Bangladesh have been blamed on logging tropical hardwoods in the
mountains of Nepal and India.
Conversely, trees can also help mitigate against drought. Like all plants,
trees release water into the atmosphere through the process of transpira-
tion. As the world’s forests shrink in total acreage, fewer greenhouse gases
such as carbon dioxide will be removed from the atmosphere, less oxygen
and water will be released into it, and the world will become a hotter, dryer
place. Scientists and policy analysts alike agree that deforestation is a ma-
jor threat to the environment. The question is whether effective policies can
be developed to reverse it or if short-term economic greed will win out
over long-term global survival.
Nancy Farm Männikkö
See also: Biodiversity; Conservation biology; Endangered plant species;

Erosion and erosion control; Forest management; Forests; Grazing and
overgrazing; Multiple-use approach; Old-growth forests; Rain forests;
Rain forests and the atmosphere; Rangeland; Reforestation; Restoration
ecology; Slash-and-burn agriculture; Soil contamination; Sustainable de-
velopment.

Bevis, William W. Borneo Log: The Struggle for Sarawak’s Forests. Seattle: Uni-
versity of Washington Press, 1995.
Colchester, Marcus, and Larry Lohmann, eds. Struggle for Land and the Fate
of the Forests. Atlantic Highlands, N.J.: Zed Books, 1993.
Dean, Warren. With Broadax and Firebrand: The Destruction of the Brazilian
Atlantic Forest. Berkeley: University of California Press, 1997.
Richards, John F., and Richard P. Tucker, eds. World Deforestation in the
Twentieth Century. Durham, N.C.: Duke University Press, 1988.
135
Deforestation
Rudel, Thomas K., and Bruce Horowitz. Tropical Deforestation: Small

Farmers and Land Clearing in the Ecuadorian Amazon. New York: Colum-
bia University Press, 1993.
Sponsel, Leslie E., Robert Converse Bailey, and Thomas N. Headland, eds.
Tropical Deforestation: The Human Dimension. New York: Columbia Uni-
Vajpeyi, Dhirendrea K., ed. Deforestation, Environment, and Sustainable De-
velopment: A Comparative Analysis. Westport, Conn.: Praeger, 2001.
Wilson, Edward O. The Future of Life. New York: Alfred A. Knopf, 2001.
136
DEMOGRAPHICS
Type of ecology: Population ecology
Demography is the study of the numbers of organisms born in a population within

a certain time period, the rate at which they survive to various ages, and the num-
ber of offspring that they produce. Many different patterns of birth, survival, and
reproduction are found among organisms in nature.
N o animal lives forever. Instead, each individual has a generalized life

history that begins with fertilization and then goes through embry-
onic development, a juvenile stage, a period in which it produces off-
spring, and finally death. There are many variations on this general theme.
Still, the life of each organism has two constants: a beginning and an
end. Many biologists are fascinated by the births and deaths of individu-
als in a population and seek to understand the processes that govern the
production of new individuals and the deaths of those already present.
The branch of biology that deals with such phenomena is called demog-
raphy.
The word “demography” is derived from Greek; demos means “peo-
ple.” For many centuries, demography was applied almost exclusively
to humans as a way of keeping written records of new births, mar-
riages, deaths, and other socially relevant information. During the first
half of the twentieth century, biologists gradually began to census pop-
ulations of naturally occurring organisms to understand their ecology
more fully. Biologists initially focused on vertebrate animals, particularly
game animals and fish. Beginning in the 1960’s and 1970’s, invertebrate an-
imals, plants, and microbes also became subjects of demographic studies.
Studies clearly show that different species of organisms vary greatly in
their demographic properties. Often, there is a clear relationship between
those demographic properties and the habitat in which these organisms
live.
Demographic Parameters
When conducting demographic studies, a demographer must gather cer-
tain types of basic information about the population. The first is the num-
ber of new organisms that appear in a given amount of time. There are two
ways that an organism can enter a population: by being born into it or by
immigrating from elsewhere. Demographers generally ignore immigra-
tion and concentrate instead on newborns. The number of new individuals
137
Demographics
born into a population during a specific time interval is termed the natality
rate. The natality rate is often based on the number of individuals already
in the population. For example, if ten newborns enter a population of a
thousand individuals during a given time period, the natality rate is 0.010.
A specific time interval must be expressed (days, months, years) for the na-
tality rate to have any meaning.
A second demographic parameter is the mortality rate, which is simply
the rate at which individuals are lost from the population by death. Losses
that result from emigration to a different population are ignored by most
demographers. Like the natality rate, the mortality rate is based on the
number of individuals in the population, and it reflects losses during a cer-
tain time period. If calculated properly, the natality and mortality rates are
directly comparable, and one can subtract the latter from the former to pro-
vide an index of the change in population size over time. The population
increases whenever natality exceeds mortality and decreases when the re-
verse is true. The absolute value of the difference denotes the rate of popu-
lation growth or decline.
When studying mortality, demographers determine the age at which
organisms die. Theoretically, each species has a natural life span that no in-
dividuals can surpass, even under the most ideal conditions. Normally,
however, few organisms reach their natural life span, because conditions
are far from ideal in nature. Juveniles, young adults, and old adults can all
die. When trying to understand the dynamics of a population, demogra-
phers therefore note whether the individuals are dying mainly as adults or
mainly as juveniles.
Patterns of Survival
Looking at it another way, demographers want to know the pattern of sur-
vival for a given population. This can best be determined by identifying a
cohort, which is defined as a group of individuals that are born at about the
same time. That cohort is then followed over time, and the number of sur-
vivors is counted at set time intervals. The census stops after the last mem-
ber of the cohort dies. The pattern of survival exhibited by the whole co-
hort is called its survivorship. Ecologists have examined the survivorship
patterns of a wide array of species, including vertebrate animals, inverte-
brates, plants, fungi, algae, and even microscopic organisms. They have
also investigated organisms from a variety of habitats, including oceans,
deserts, rain forests, mountain peaks, meadows, and ponds. Survivorship
patterns vary tremendously.
Some species have a survivorship pattern in which the young and
middle-aged individuals have a high rate of survival, but old individu-
138
Demographics
als die in large numbers. Several species of organisms that live in nature,
such as mountain sheep and rotifers (tiny aquatic invertebrates), ex-
hibit this survivorship pattern. At the other extreme, many species ex-
hibit a survivorship pattern in which mortality is heaviest among the
young. Those few individuals that are fortunate enough to survive the
period of heavy mortality then enjoy a high probability of surviving until
the end of their natural life span. Examples of species that have this pat-
tern include marine invertebrates such as sponges and clams, most spe-
cies of fish, and parasitic worms. An intermediate pattern is also observed,
in which the probability of dying stays relatively constant as the cohort
gets older. American robins, gray squirrels, and hydras all display this pat-
tern.
These survivorship patterns are usually depicted on a graph that has
the age of individuals in the cohort on the x-axis and the number of survi-
vors on the y-axis. Each of the three survivorship patterns gives a different
curve when the number of survivors is plotted as a function of age. In the
first pattern (high survival among juveniles), the curve is horizontal at first
but then swings downward at the right of the graph. In the second pattern
(low survival among juveniles), the curve drops at the left of the graph but
then levels out to form a horizontal line. The third survivorship pattern
(constant mortality throughout the life of the cohort) gives a straight line
that runs from the upper-left corner of the graph to the lower right (this is
best seen when the y-axis is expressed as the logarithm of the number of
survivors).
In the first half of the twentieth century, demographers Raymond Pearl
and Edward S. Deevey labeled each survivorship pattern: Type I is high
survival among juveniles, type II is constant mortality through the life of
the cohort, and type III is low survival among juveniles. That terminology
became well entrenched in the biological literature by the 1950’s. Few spe-
cies exhibit a pure type I, II, or III pattern, however; instead, survivorship
varies so that the pattern may be one type at one part of the cohort’s exis-
tence and another type later. Perhaps the most common survivorship pat-
tern, especially among vertebrates, is composed of a type III pattern for ju-
veniles and young adults followed by a type I pattern for older adults. This
pattern can be explained biologically. Most species tend to suffer heavy ju-
venile mortality because of predation, starvation, cannibalism, or the in-
ability to cope with a stressful environment. Juveniles that survive this
hazardous period then become strong adults that enjoy relatively low mor-
tality. As time passes, the adults reach old age and ultimately fall victim to
disease, predation, and organ-system failure, thus causing a second down-
ward plunge in the survivorship curve.
139
Demographics
Patterns of Reproduction
Demographers are not interested only in measuring the survivorship of co-
horts. They also want to understand the patterns of reproduction, espe-
cially among females. Different species show widely varying patterns of
reproduction. For example, some species, such as octopuses and certain
salmon, reproduce only once in their life and then die soon afterward. Oth-
ers, such as humans and most birds, reproduce several or many times in
their lives. Species that reproduce only once accumulate energy through-
out their lives and essentially put all of it into producing young. Reproduc-
tion essentially exhausts them to death. Conversely, those that reproduce
several times devote only a small amount of their energy to each reproduc-
tive event.
Species also vary in their fecundity, which is the number of offspring
that an individual makes when it reproduces. Large mammals have low fe-
cundity, because they produce only one or two progeny at a time. Birds,
reptiles, and small mammals have higher fecundity because they typically
produce a clutch or litter of several offspring. Fish, frogs, and parasitic
worms have very high fecundity, producing hundreds or thousands of off-
spring.
A species’ pattern of reproduction is often related to its survivorship.
For example, a species with low fecundity or one that reproduces only
once tends to have type I or type II survivorship. Conversely, a species that
produces huge numbers of offspring generally shows type III survivor-
ship. Many biologists are fascinated by this interrelationship between
survivorship and reproduction. Beginning in the 1950’s, some demogra-
phers proposed mathematically based explanations as to how the interre-
lationship might have evolved as well as the ecological conditions in
which various life histories would be expected. For example, some demog-
raphers predicted that species with low fecundity and type I survival
should be found in undisturbed, densely populated areas (such as a tropi-
cal rain forest). In contrast, species with high fecundity and type III sur-
vival should prevail in places that are either uncrowded or highly dis-
turbed (such as an abandoned farm field). Ecologists have conducted field
studies of both plants and animals to determine whether the patterns that
actually occur in nature fit the theoretical predictions. In some cases the
predictions were upheld, but in others they were found to be wrong and
had to be modified.
Age Structures and Sex Ratios

Another feature of a population is its age structure, which is simply the
number of individuals of each age. Some populations have an age struc-
140
Demographics
ture characterized by many juveniles and only a few adults. Two situations
could account for such a pattern. First, the population could be rapidly ex-
panding, with the adults successfully reproducing many progeny that are
enjoying high survival. Second, the population could be producing many
offspring that have type III survival. In this second case, the size of the pop-
ulation can remain constant or even decline. Other populations have a dif-
ferent age structure, in which the number of juveniles only slightly exceeds
the number of adults. Those populations tend to remain relatively constant
over time. Still other populations have an age structure in which there are
relatively few juveniles and many adults. Those populations are probably
declining or are about to decline because the adults are not successfully re-
producing.
Since most animals are unisexual, an important demographic character-
istic of a population is its sex ratio, defined as the ratio of males to females.
While the ratio for birds and mammals tends to be 1:1 at conception (the
fertilization of an egg), it tends to be weighted toward males at birth, be-
cause female embryos are slightly less viable. After birth, the sex ratio for
mammals tends to favor females, because young males suffer higher mor-
tality. The posthatching ratio in birds tends to remain skewed toward
males, because females devote considerable energy to producing young
and suffer higher mortality. As a result, male birds must compete with one
another for the opportunity to mate with the scarcer females.
The Age-Specific Approach

To understand the demography of a particular species, one must collect in-
formation about its survivorship and reproduction. The best survivorship
data are obtained when a demographer follows a group of newly born
organisms (this being a cohort) over time, periodically counting the survi-
vors until the last one dies. Although that sounds relatively straightfor-
ward, many factors complicate the collection of survivorship data; demog-
raphers must be willing to adjust their methods to fit the particular species
and environmental conditions.
First, a demographer must decide how many newborns should be in-
cluded in the cohort. Survivorship is usually based on one thousand new-
borns, but few studies follow that exact number. Instead, demographers
follow a certain number of newborns and multiply or divide their data so
that the cohort is expressed as one thousand newborns. For example, one
may choose to follow five hundred newborns; the number of survivors is
then multiplied by two. Demographers generally consider cohorts com-
posed of fewer than one hundred newborns to be too small. Second, meth-
ods of determining survivorship are much more different for highly motile
141
Demographics
organisms, such as mammals and birds, than for more sedentary ones,
such as bivalves (oysters and clams). To determine survivorship of a sed-
entary species, demographers often find some newborns during an initial
visit to a site and then periodically revisit that site to count the number of
survivors. Highly motile animals are much more difficult to census because
they do not stay in one place waiting to be counted. Vertebrates and large
invertebrates can be tagged, and individuals can be followed by subse-
quently recapturing them. Some biologists use small radio transmitters to
follow highly active species. The demography of small invertebrates such
as insects is best determined when there is only one generation per year
and members of the population are all of the same age-class. For such spe-
cies, demographers merely count the number present at periodic intervals.
Third, the frequency of the census periods varies from species to spe-
cies. Short-lived species, such as insects, must be censused every week or
two. Longer-lived species need to be counted only once a year. Fourth, the
definition of a “newborn” may be troublesome, especially for species with
complex life cycles. Demographic studies usually begin with the birth of
an infant. Some would argue, however, that the fetus should be included in
the analysis because the starting point is really conception. Many seden-
tary marine invertebrates (sponges, starfish, and barnacles) have highly
motile larval stages, and these should be included in the analysis for
survivorship to be completely understood. Parasitic roundworms and
flatworms that have numerous juvenile stages, each found inside a differ-
ent host, are particularly challenging to the demographer.
The Time-Specific Approach

The survivorship of long-lived species, such as large mammals, is really
impossible to determine by the methods given above. Because of their
sheer longevity, one could not expect a scientist to be willing to wait de-
cades or centuries until the last member of a cohort dies. Demographers at-
tempt to overcome this problem by using the age distribution of organisms
that are alive at one time to infer cohort survivorship. This is often termed a
“horizontal” or “time-specific” approach, as opposed to the “vertical” or
“age-specific” approach that requires repeated observations of a single co-
hort. For example, one might construct a time-specific survivorship curve
for a population of fish by live-trapping a sufficiently large sample, count-
ing the rings on the scales on each individual (which for many species is
correlated with the age in years), and then determining the number of one-
year-olds, two-year-olds, and so on. Typically, demographers who use age
distributions to infer age-specific survivorship automatically assume that
natality and mortality remain constant from year to year. That is often not
142
Demographics
the case, however, because environmental conditions often change over

time. Thus, demographers must be cautious when using age distribution
data to infer survivorship.
Methods for determining fecundity are relatively straightforward.
Typically, fertile individuals are collected, their ages are determined, and
the number of progeny (eggs or live young) are counted. Species that re-
produce continually (parasitic worms) or those that reproduce several
times a year (small mammals and many insects) must be observed over a
period of time.
Demographers usually want to determine whether the production of
new offspring (natality) balances the losses attributable to mortality. To ac-
complish this, they construct a life table, which is a chart with several col-
umns and rows. Each row represents a different age of the cohort, from
birth to death. The columns show the survival and fecundity of the cohort.
By recalculating the survivorship and fecundity information, demogra-
phers can compute several interesting aspects of the cohort, including the
life expectancy of individuals at different ages, the cohort’s reproductive
value (which is the number of progeny that an individual can expect to
produce in the future), the length of a generation for that species, and the
growth rate for the population.
Uses of Demography
Demographic techniques have been applied to nonhuman species, partic-
ularly by wildlife managers, foresters, and ecologists. Wildlife managers
seek to understand how a population is surviving and reproducing within
a certain area, and therefore to determine whether it is increasing or de-
creasing over time. With that information, a wildlife biologist can then esti-
mate the effect of hunting or other management practice on the popula-
tion. By extension, fisheries biologists can also make use of demographic
techniques to determine the growth rate of the species of interest. If the
population is determined to be increasing, it can be harvested without fear
of depleting the population. Alternatively, one can conduct demographic
analyses to see whether certain species are being overfished.
An often unappreciated benefit of survivorship analyses is that they can
help ecologists pinpoint factors that limit population growth in an area.
This may be especially important in efforts to prevent rare animals and
plants from becoming extinct. Once the factor is identified, the population
can be appropriately managed. Increasing amounts of public and private
money are allocated each year to biologists who conduct demographic
studies on rare species.
Kenneth M. Klemow
143
Demographics
See also: Adaptive radiation; Biodiversity; Biogeography; Clines, hybrid

zones, and introgression; Convergence and divergence; Extinctions and
evolutionary explosions; Gene flow; Genetic diversity; Genetic drift; Hu-
man population growth; Insect societies; Nonrandom mating, genetic
drift, and mutation; Population analysis; Population fluctuations; Popula-
tion genetics; Population growth; Reproductive strategies.

uals, Populations, and Communities. 3d ed. Boston: Blackwell, 1996.
Begon, Michael, Martin Mortimer, and David J. Thompson. Population
Ecology: A Unified Study of Animals and Plants. 3d ed. Cambridge, Mass.:
Blackwell, 1996.
Brewer, Richard. The Science of Ecology. 2d ed. Fort Worth, Tex.: Saunders
College Publishing, 1994.
Elseth, Gerald D., and Kandy D. Baumgardner. Population Biology. New
York: Van Nostrand, 1981.
Gotelli, Nicholas J. A Primer of Ecology. 2d ed. Sunderland, Mass.: Sinauer
Associates, 1998.
Hutchinson, G. Evelyn. An Introduction to Population Ecology. New Haven,
Conn.: Yale University Press, 1978.
Smith, Robert Leo. Elements of Ecology. 4th ed. San Francisco, Calif.:
Benjamin/Cummings, 2000.
Wilson, Edward O., and William Bossert. A Primer of Population Biology.
Sunderland, Mass.: Sinauer Associates, 1977.
144
DENDROCHRONOLOGY
Dendrochronology is the science of examining and comparing growth rings in

both living and aged woods to draw inferences about past ecosystems and environ-
mental conditions.
I n forested regions with seasonal climates, trees produce a growth ring to

correspond with each growing season. At the beginning of the growing
season, when conditions are optimum, the vascular cambium produces
many files of large xylem cells that form wood. As the conditions become
less optimal, the size and number of cells produced decreases until growth
stops at the end of the growing season. These seasonal differences in size
and number of cells produced are usually visible to the unaided eye. The
layers produced during rapid early growth appear relatively light-colored
because the volume of the large cells is primarily intracellular space. These
layers are frequently called springwood because in northern temperate re-
gions spring is the beginning of the growing season. Wood formed later,
summerwood, is darker because the cells are smaller and more tightly
compacted. The juxtaposition of dark summerwood of one year with the
light springwood of the following year marks a distinct line between
growth increments. The width of the ring between one line and the next
measures the growth increment for a single growing season. If there is a
single growing season per year, as in much of the temperate world, then a
tree will produce a single annual ring each year.
Tree Rings and Climate

Leonardo da Vinci is credited with counting tree rings in the early 1500’s to
determine “the nature of past seasons,” but it was not until the early 1900’s
that dendrochronology was established as a science. Andrew Douglass, an
astronomer interested in relating sunspot activity to climate patterns on
Earth, began to record the sequences of wide and narrow rings in the wood
of Douglas firs and ponderosa pines in the American Southwest. Ori-
ginally, trees were cut down in order to examine the ring patterns, but in
the 1920’s Douglass began to use a Swedish increment borer to remove
core samples from living trees. This instrument works like a hollow drill
that is screwed into a tree by hand. When the borer reaches the center of the
tree it is unscrewed, and the wood core sample inside is withdrawn with
the borer. The small hole quickly fills with sap, and the tree is unharmed.
145
Dendrochronology
Borers range in size from 20 centimeters to 100 centimeters or more in

length, so with care, samples can be taken from very large, very old living
trees.
Counting backward in the rings is counting backward in time. By corre-
lating the size of a ring with the known regional climate of the year the ring
was produced, a researcher can calibrate a core sample to indicate the sur-
rounding climate during any year of the tree’s growth. By extending his
work to sequoias in California, Douglass was able to map a chronology ex-
tending back three thousand years.
Tree Rings and History

In order to extend his chronologies so far back in time, Douglass devised
the method of cross-dating. By matching distinctive synchronous ring pat-
terns from living and dead trees of the same species in a region, researchers
can extend the pattern further into the past than the lifetime of the younger
tree. Archaeologists quickly realized that this was a tool that could help to
assign the age of prehistoric sites by determining the age of wood artefacts
and construction timbers. In this way archaeologists could calculate the
age of pre-Columbian southwestern ruins, such as the cliff dwellings at
Mesa Verde, Arizona, by cross-dating living trees with dead trees and the
latter with timbers from the sites. In 1937 Douglass established the Labora-
tory of Tree-Ring Research at the University of Arizona, which continues to
be a major center of dendrochronological research.
Fine-Tuning
In the mid-1950’s Edmund Schulman confirmed the great age of living
bristlecone pines in the Inyo National Forest of the White Mountains of
California. In 1957 he discovered the Methuselah Tree, which was more
than forty-six hundred years old. The section of forest in which he worked
is now known as the Ancient Bristlecone Pine Forest. During the next
thirty years, Charles Ferguson extended the bristlecone chronology of this
area back 8,686 years. This sequence formed the basis for calibrating the
technique of radiocarbon dating. In the 1960’s, radiocarbon analysis began
to be used to determine the age of organic (carbon-based) artefacts from
ancient sites. It has the advantage of being applicable to any item made of
organic material but the disadvantage of having a built-in uncertainty of
2 percent or more. Tree-ring chronologies provide an absolute date against
which radiocarbon analyses of wood samples from a site can be compared.
At about the same time, Valmore LaMarche, a young geologist, began to
study root growth of the ancient trees to determine how they could be used
to predict the erosional history of a site. By cross-referencing growth ring
146
Dendrochronology
Dendrochronology, or tree-ring counting, can be used to assess the age of a tree be-
cause the width of the ring between one line and the next measures the growth in-
crement for a single growing season. (PhotoDisc)
asymmetry to degree of exposure and slope profiles, he was able to esti-

mate rates of soil erosion and rock weathering, which in turn could be
cross-referenced to the climatic conditions predicted by growth rings in the
stem. LaMarche and his colleagues, particularly Harold Fritts, continued
to “fine-tune” the reading of growth rings to be able to take into account
factors such as soil characteristics, frost patterns, and daily, weekly, and
monthly patterns.
The Oldest Tree

The Methuselah Tree, mentioned above, is the oldest known living tree.
Schulman also cored a 4,700-year-old living specimen in the White Moun-
tains, but he did not name it or identify its location. While most of the liv-
ing specimens older than 4,000 years are found in the White Mountains,
the oldest living tree was discovered in the Wheeler Peak area of what is
now Great Basin National Park in eastern Nevada. This tree, variously
known as WPN-114 and the Prometheus Tree, was estimated to be between
4,900 and 5,100 years old when it was cut down in 1964 as part of a research
project. The controversy that followed has left many interesting but unan-
swered questions.
147
Dendrochronology
See also: Evolution of plants and climates; Forests; Global warming; Old-
growth forests; Paleoecology.

Cohen, Michael P. Garden of Bristlecones: Tales of Change in the Great Basin.
Reno: University of Nevada Press, 2000.
Cook, E. R., and L. A. Kairiustis, eds. Methods of Dendrochronology: Applica-
tions in the Environmental Sciences. Boston: Kluwer Academic, 1987.
Harlow, William M. Inside Wood: Masterpiece of Nature. Washington, D.C.:
The American Forestry Association, 1970.
McGraw, Donald J. Andrew Ellicott Douglass and the Role of the Giant Sequoia
in the Development of Dendrochronology. Lewiston, N.Y.: Edwin Mellen
Press, 2001.
Stokes, Marvin A., and Terah L. Smiley. An Introduction to Tree-Ring Dating.
Tucson: University of Arizona Press, 1996.
148
DESERTIFICATION
Type of ecology: Ecosystem ecology
Desertification is the degradation of arid, semiarid, and dry, subhumid lands as a

result of human activities or climatic variations, such as a prolonged drought.
D esertification is recognized by scientists and policymakers as a major

economic, social, and environmental problem in more than one hun-
dred countries. It impacts about one billion people throughout the world.
Deserts are climatic regions that receive fewer than 25 centimeters (10
inches) of precipitation per year. They constitute the most widespread of
all climates of the world, occupying 25 percent of the earth’s land area.
Most deserts are surrounded by semiarid climates referred to as steppes,
which occupy 8 percent of the world’s lands. Deserts occur in the interior
of continents, on the leeward side of mountains, and along the west sides
of continents in subtropical regions. All the world’s deserts risk further de-
sertification.
Deserts of the World

The largest deserts are in North Africa, Asia, Australia, and North Amer-
ica. Four thousand to six thousand years ago, these desert areas were less
extensive and were occupied by prairie or savanna grasslands. Rock paint-
ings found in the Sahara Desert show that humans during this era hunted
buffalo and raised cattle on grasslands where giraffes browsed. The region
near the Tigris and Euphrates Rivers in the Middle East was also fertile. In
the desert of northwest India, cattle and goats were grazed, and people
lived in cities that have long since been abandoned. The deserts in the
southwestern region of North America appear to have been wetter, accord-
ing to the study of tree rings (dendrochronology) from that area. Ancient
Palestine, which includes the Negev Desert of present-day Israel, was lush
and was occupied by three million people.
Scientists use various methods to determine the historical climatic con-
ditions of a region. These methods include studies of the historical distri-
bution of trees and shrubs determined by the deposit patterns in lakes and
bogs, patterns of ancient sand dunes, changes in lake levels through time,
archaeological records, and tree rings.
The earth’s creeping deserts supported approximately 720 million
people, or one-sixth of the world’s population, in the late 1970’s. Accord-
149
Desertification
ing to the United Nations, the world’s hyperarid or extreme deserts are
the Atacama and Peruvian Deserts (located along the west coast of South
America), the Sonoran Desert of North America, the Takla Makan Desert
of Central Asia, the Arabian Desert of Saudi Arabia, and the Sahara
Desert of North Africa, which is the largest desert in the world. The arid
zones surround the extreme desert zones, and the semiarid zones sur-
round the arid zones. Areas that surround the semiarid zones have a high
risk of becoming desert. By the late 1980’s the expanding deserts were
claiming about 15 million acres of land per year, or an area approximately
the size of the state of West Virginia. The total area threatened by desertifi-
cation equaled about 37.5 million square kilometers (14.5 million square
miles).
Causes of Desertification
Desertification results from a two-prong process: climatic variations and
human activities. First, the major deserts of the world are located in areas
of high atmospheric pressure, which experience subsiding dry air unfavor-
able to precipitation. Subtropical deserts have been experiencing pro-
longed periods of drought since the late 1960’s, which causes these areas to
be dryer than usual.
The problem of desertification was identified in the late 1960’s and early
1970’s as a result of severe drought in the Sahel Desert, which extends
along the southern margin of the Sahara in West Africa. Rainfall has de-
clined an average of 30 percent in the Sahel. One set of scientific studies of
the drought focuses on changes in heat distribution in the ocean. A correla-
tion has been found between sea surface temperatures and the reduction of
rainfall in the Sahel. The Atlantic Ocean’s higher surface temperatures
south of the equator and lower temperatures north of the equator west of
Africa are associated with lower precipitation in northern tropical Africa.
However, the cause for the change in sea surface temperature patterns has
not been determined.
Another set of studies is associated with land-cover changes. Lack of
rain causes the ground and soils to get extremely dry. Without vegetative
cover to hold it in place, thin soil blows away. As the water table drops
from the lack of the natural recharge of the aquifers and the withdrawal of
water by desert dwellers, inhabitants are forced to migrate to the grass-
lands and forests at fringes of the desert. Overgrazing, overcultivation, de-
forestation, and poor irrigation practices (which can cause salinization of
soils) eventually lead to a repetition of the process, and the desert begins to
encroach. These causes are influenced by changes in population, climate,
and social and economic conditions.
150
Desertification
Desertification of Africa
Mediterranean Sea
Tropic of Cancer
Gulf of Guinea
Equator
Indian
Ocean
True desert Atlantic
Acute risk of Ocean
desertification
Moderate to
great risk
Tropic of Capricorn
The fundamental cause of desertification, therefore, is human activity.

This is especially true when environmental stress occurs because of sea-
sonal dryness, drought, or high winds. Many different forms of social, eco-
nomic, and political pressure cause the overutilization of these dry lands.
People may be pushed onto unsuitable agricultural land because of land
shortages, poverty, and other forces, while farmers overcultivate the fields
in the few remaining fertile land areas.
Atmospheric Consequences
A reduction in vegetation cover and soil quality may impact the local cli-
mate by causing a rise in temperatures and a reduction in moisture. This
can, in turn, impact the area beyond the desert by causing changes in the
151
Desertification
climate and atmospheric patterns of the region. It is predicted that by the

year 2050 substantial changes in vegetation cover in humid and subhumid
areas will occur and cause substantial regional climatic changes. Desertifi-
cation is a global problem because it causes the loss of biodiversity as well
as the pollution of rivers, lakes, and oceans. As a result of excessive rainfall
and flooding in subhumid areas, fields lacking sufficient vegetation may
be eroded by runoff.
Greenhouse Effect
Desertification and even the efforts to combat it may be impacting climatic
change because of the emission and absorption of greenhouse gases. The
decline in vegetation and soil quality can result in the release of carbon,
while revegetation can influence the absorption of carbon from the atmo-
sphere. The use of fertilizer to reclaim dry lands may cause an increase in
nitrous oxide emissions. Although scientists involved in studies of rising
greenhouse gases have not been able to gather evidence conclusive enough
to support such theories, evidence of the impact of greenhouse gases on
global warming continues to accumulate.
Policy Actions
As a result of the Sahelian drought, which lasted from 1968 to 1973, repre-
sentatives from various countries met in Nairobi, Kenya, in 1977 for a
United Nations conference on desertification. The conference resulted in
the Plan of Action to Combat Desertification. The plan listed twenty-eight
measures to combat land degradation by national, regional, and interna-
tional organizations. A lack of adequate funding and commitment by gov-
ernments caused the plan to fail. When the plan was assessed by the
United Nations Environment Programme (UNEP), it found that little had
been accomplished and that desertification had increased.
As a result of the 1977 United Nations conference, several countries de-
veloped national plans to combat desertification. One example is Kenya,
where local organizations have worked with primary schools to plant five
thousand to ten thousand seedlings per year. One U.S.-based organization
promotes reforestation by providing materials to establish nurseries, train-
ing programs, and extension services. Community efforts to combat deser-
tification have been more successful, and UNEP has recognized that such
projects have a greater success rate than top-down projects. The Earth
Summit, held in Rio de Janeiro, Brazil, in 1992, supported the concept of
sustainable development at the community level to combat the problem of
desertification.
Roberto Garza
152
Desertification
See also: Biomes: determinants; Biomes: types; Deserts; Ecosystems: defi-

nition and history; Erosion and erosion control; Global warming; Green-
house effect; Hydrologic cycle; Rain forests and the atmosphere; Soil; Soil
contamination.

Bryson, Reid A., and Thomas J. Murray. Climates of Hunger: Mankind and
the World’s Changing Weather. Madison: University of Wisconsin Press,
1977.
Glantz, Michael H., ed. Desertification: Environmental Degradation in and
Around Arid Lands. Boulder, Colo.: Westview Press, 1977.
Grainger, Alan. Desertification: How People Make Deserts, Why People Can
Stop, and Why They Don’t. London: International Institute for Environ-
ment and Development, 1982.
Hulme, Mike, and Mick Kelly. “Exploring the Links Between Desertifica-
tion and Climate Change.” Environment, July/August, 1993.
Mainguet, Monique. Aridity: Droughts and Human Development. New York:
Springer, 1999.
_______. Desertification: Natural Background and Human Mismanagement. 2d
ed. New York: Springer-Verlag, 1994.
Matthews, Samuel W. “What’s Happening to Our Climate.” National Geo-
graphic, November, 1976.
Postel, Sandra. “Land’s End.” Worldwatch, May/June, 1989.
153
DESERTS
Regions characterized by 10 inches or less of precipitation per year are considered

deserts. Desert ecosystems are subject to disruption by human activities.
D eserts are regions, or biomes, too dry to support grasslands or forest

vegetation but with enough moisture to allow specially adapted
plants to live. In deserts, hot days alternate with cold nights. Ninety per-
cent of incoming solar radiation reaches the ground during the day, and 90
percent of that is radiated back out into space at night, the result of the ab-
sence of clouds, low humidity, and sparse vegetation. The surface of the
ground in desert areas is devoid of a continuous layer of plant litter and is
usually rocky or sandy. Nutrient cycling in deserts is tight, with phospho-
rus and nitrogen typically in short supply.
How Deserts Form

Most large landmasses have interior desert regions. Air masses blown in-
land from coastal areas lose their moisture before reaching the interior. Ex-
amples include the Gobi Desert in Mongolia and parts of the Sahara Desert
in Africa.
Another factor in the formation of deserts is the rain-shadow effect. If
moisture-laden air masses bump up against a mountain range, the air mass
is deflected upward. As the air mass rises, it cools, and moisture precipi-
tates as rain or snow on the windward side of the mountain range. As the
air mass passes over the mountain range, it begins to descend. Because it
lost most of its moisture on the windward side, the air mass is dry. As it de-
scends, the air heats up, creating drier conditions on the leeward side of the
mountain range. Sometimes these differences in moisture are so pro-
nounced that different plant communities grow on the windward and lee-
ward sides.
Latitude can also influence desert formation. Most deserts lie between
15 and 35 degrees north or south latitude. At the equator, the sun’s rays hit
the earth straight on. Moist equatorial air, warmed by intense heat from the
sun, rises. As this air rises, it cools and loses its moisture, which falls as
rain; this is why it usually rains every day in the equatorial rain forests. The
Coriolis force causes the air masses to veer off, to the north in the Northern
Hemisphere and to the south in the Southern Hemisphere. The now-dry
air begins to descend and warm, reaching the ground between 15 and 35
154
Deserts
degrees north and south latitude, creating the belt of deserts circling the
globe between these latitudes.
Deserts can also form along coastlines next to cold-water ocean cur-
rents, which chill the air above them, decreasing their moisture content.
Offshore winds blow the air above cold ocean waters back out to sea. In
deserts, rain is infrequent, creating great hardships for the native plants
and animals. The main source of moisture for the plants and animals of
coastal deserts is fog.
Types of Deserts
Depending upon whether the precipitation comes from rain or snow,
deserts can be divided into hot (rain) or cold (snow) deserts. The deserts of
Arabia, Australia, Chihuahua, Kalahari, Monte, Sonora, and Thar are all
considered hot deserts, found in lower latitudes. Cold deserts, found at
higher latitudes, include the Atacama, Gobi, Basin, Iranian, Namib, and
Turkestan deserts. Regardless of whether the desert is hot or cold, organ-
isms living within desert biomes have to adapt to cope with the scarcity of
water and violent swings of temperature. North America contains four dif-
ferent deserts that are usually defined by their characteristic vegetation,
which ecologists call indicator species.
Although desert climates are characterized by less than 10 inches of precipitation

per year, they can support a broad variety of plant and animal life adapted to sur-
vive such arid climates. Cacti, for example, have developed a method of photosyn-
thesis that maximizes water retention, as well as protective spines that inhibit
predators that might feed on them. (PhotoDisc)
155
Deserts
In Mexico’s Chihuahuan Desert, lechuguilla (Agave lechuguilla) is the

indicator species. Fibers from lechuguilla can be made into nets, bas-
kets, mats, ropes, and sandals. Its stems yield a soap substitute, and its
pulp has been used as a spot remover. Certain compounds in lechuguilla
are poisonous and were once used to poison the tips of arrows and as
fish poisons. Two of the most common plants in the Chihuahuan Desert are
creosote bush (Larrea divaricata) and soaptree yucca (Yucca elata). Cacti in
this desert are numerous and diverse, especially the prickly pears and
chollas.
The Joshua tree (Yucca brevifolia), is the indicator species of the Mojave
Desert in Southern California. Nearly one-fourth of all the Mojave Desert
plants are endemics, including the Joshua tree, Parry saltbush, Mojave
sage, and woolly bur sage.
The Great Basin Desert, situated between the Sierra Nevada and the
Rocky Mountains, is a cold desert, with fewer plant species than other
North American deserts. Great Basin Desert plants are small to medium-
size shrubs, usually sagebrushes or saltbushes. The indicator species of the
Great Basin is big sagebrush (Artemisia tridentata). Other common plants
are littleleaf horsebrush and Mormon tea. The major cactus species is the
Plains prickly pear.
In the Sonoran Desert in Mexico, California, and Arizona, plants come
in more shapes and sizes than in the other North American deserts, espe-
cially in the Cactaceae. The indicator species of the Sonoran Desert is the sa-
guaro cactus (Carnegiea gigantecus).
The Sahara Desert of northern Africa is the world’s largest, at 3.5 mil-
lion square miles. The Northern Hemisphere also contains the Arabian,
Indian, and Iranian deserts and the Eurasian deserts: the Takla Makan, Tur-
kestan, and Gobi. Deserts in the Southern Hemisphere include the Austra-
lian, Kalahari, Namib, Atacama-Peruvian (the world’s driest), and the Pata-
gonian.
Desert Vegetation
Many typical desert perennial plants, such as members of the Cactaceae (the
cactus family), have thick, fleshy stems or leaves with heavy cuticles,
sunken stomata (pores), and spiny defenses against browsing animals. The
spines also trap a layer of air around the plant, retarding moisture loss.
Desert plants, many of which photosynthesize using C4 or CAM (cras-
sulean acid metabolism), live spaced out from other plants. Many desert
plants are tall and thin, to minimize the surface area exposed to the stron-
gest light. For example, the entire stem of the Saguaro cactus is exposed to
sunlight in the early morning and late afternoon; at noon, only the tops
156
Deserts
of the stems receive full sun. These traits allow the plants to cope with
heat stress and competition for water and avoid damage from herbivores
(plant-eating animals).
Where the mixture of heat and water stress is less severe, perennial
bushes of the Chenopodiaceae (goosefoot family) or Asteraceae (sunflower
family) form clumps of vegetation surrounded by bare ground. Numerous
annuals, called ephemerals, can grow prolifically, if only briefly, following
rainfall.
Unrelated plant families from different desert areas of the world show
similar adaptations to desert conditions. This has resulted from a process
called convergent evolution.
Animal Life in Deserts

Animals living in the desert include mammals, birds and fish, reptiles and
amphibians, and insects and spiders. A few examples of mammals include
bats, bighorn sheep, bobcats, and coyotes. Desert bats, members of the
suborder Microchioptera, are often considered to be flying mice, but they
are more closely related to primates. Bats are unique among mammals be-
cause they can fly. Most bat species also possess a system of acoustic orien-
tation, technically known as echolocation. The bighorn sheep lives in dry,
desert mountain ranges and foothills near rocky cliffs. Its body is com-
pact and muscular; the muzzle is narrow and pointed; the ears, short
and pointed; the tail, very short. The fur is deerlike and usually a shade
of brown, with whitish rump patches. Bighorns are grazers, consuming
grasses, sedges, and other low-lying plants. The bobcat (Felis rufus) has
long legs, large paws, and a short tail (six to seven inches long), with aver-
age body weight of fifteen to twenty pounds. However, it is quite fierce
and is equipped to kill animals as large as deer. The desert coyote (Canis
latrans), a member of the dog family, weighs about twenty pounds, less
than half the weight of its mountain kin, which can weigh up to fifty
pounds. Its body color is light gray or tan, which helps it to reflect heat and
blend in in the desert.
A variety of birds reside in the desert due to the abundance of insects
and spiders. Golden eagles (Aquila chrysaetos) get their name from the
golden feathers on the back of their necks. They are birds of the open coun-
try, building large stick nests in trees or cliff walls where they have plenty
of room to maneuver. Adults weigh 9 to 12.5 pounds, with females usually
larger than males. Ravens (Corvus corax) are the largest birds of the crow
family, averaging twenty-four inches tall, with a wingspan of forty-six to
fifty-six inches. Ravens are strong fliers that can soar like a hawk, and they
may form large flocks of over several hundred individuals during their au-
157
Deserts
tumn migration. The American turkey (Meleagris gallopavo), the largest up-
land game bird in North America, is thirty-six to forty-eight inches long,
with a four- to five-foot wingspan. Males average ten inches longer than fe-
males, which are paler and of a more buff color. Turkeys inhabit a variety of
habitats from open grassland and fields to open woodlands and mature
deciduous or coniferous forests.
Many species of reptiles and amphibians live in the desert, including
the black-collared lizard (genus Crotaphytus), bullfrog (Rana catesbeiana),
desert dinosaur (orders Saurischia and Ornithischia), desert iguana (Dispso-
saurus dorsalis), rattlesnake (genus Crotalus), and many others. Insects
and spiders include dragonflies (suborder Anisoptera), scorpions (order
Scorpionida), and black widow spiders (Latrodectus hesperus). It is nothing
short of a miracle that such an abundance of life, both plant and animal,
can survive in the extreme conditions of the desert.
Animal Survival in the Desert

Among the thousands of desert animal species, many have remarkable be-
havioral and structural adaptations for avoiding excess heat. Equally inge-
nious are the diverse mechanisms various animal species have developed
to acquire, conserve, recycle, and actually manufacture water.
Certain species of birds, such as the Phanopepla, breed during the rela-
tively cool spring, then leave the desert for cooler areas at higher elevations
or along the Pacific coast. The Costa’s hummingbird begins breeding in
late winter and leaves in late spring when temperatures become extreme.
Many birds, as well as other mammals and reptiles, are crepuscular, mean-
ing they are active only at dusk and again at dawn. Many animals, includ-
ing bats, many snakes, most rodents, and some larger animals such as
foxes and skunks, are nocturnal, restricting all their activities to the cooler
temperatures of the night and sleeping in a cool den, cave, or burrow by
day. A few desert animals, such as the round-tailed ground squirrel, sleep
away the hottest part of summer and also hibernate in winter to avoid the
cold season. Yet other animals, such as desert toads, remain dormant deep
in the ground until the summer rains fill ponds. They then emerge, breed,
lay eggs, and replenish their body reserves of food and water for another
long period.
Various mechanisms are employed to dissipate heat absorbed by desert
animals. Many mammals have long appendages to release body heat into
their environment. The enormous ears of jackrabbits, with their many
blood vessels, dissipate heat when the animal is resting in a cool, shady lo-
cation. Their close relatives in cooler regions have much shorter ears. New
World vultures, dark in color and thus absorbing considerable heat in the
158
Deserts
desert, excrete urine on their legs to cool them by evaporation, and circu-
late the cooled blood back through the body. Many desert animals are paler
than their relatives elsewhere, ensuring that they not only suffer less heat
absorption, but also are less conspicuous to predators in the bright, pallid
surroundings.
The mechanisms by which water is retained by desert animals are even
more elaborate. Reptiles and birds excrete metabolic wastes in the form of
uric acid, an insoluble white compound, wasting very little water in the
process. Other animals retain water by burrowing into moist soil during
the dry daylight hours. Some predatory and scavenging animals can ob-
tain their entire water needs from the food they eat. Most mammals, how-
ever, need access to a good supply of fresh water at least every few days, if
not daily, due to the considerable water loss from excretion of urea, a solu-
ble compound.
Many desert animals obtain water from plants, particularly succulent
ones such as cactus and saguaro. Many species of insects thrive in the
desert, as they tap plant fluids for water and nectar. The abundance of in-
sect life permits insectivorous birds, bats, and lizards to thrive in the
desert. Certain desert animals, such as kangaroo rats, have multiple adap-
tation mechanisms to acquire and conserve water. First, they live in under-
ground dens that they seal off to block out heat and to recycle the moisture
from their own breathing. Second, they have specialized kidneys with ex-
tra microscopic projections to extract most of the water from their urine
and return it to the bloodstream. Third, and most fascinating of all, they ac-
tually manufacture their water metabolically from the digestion of dry
seeds. These are just a few examples of the variety of ingenious adapta-
tions animals use to survive in the desert, overcoming the extremes of heat
and the paucity of water.
Habitat Loss
Urban and suburban sprawl paves over desert land and destroys habitat
for plants and animals, some of which are endemic to specific deserts.
Farmers and metropolitan-area builders can tap into critical desert water
supplies, changing the hydrology of desert regions. Off-road vehicles can
destroy plant and animal life and leave tracks that may last for decades.
Carol S. Radford and Yujia Weng
See also: Biomes: determinants; Biomes: types; Desertification; Ecosys-

tems: definition and history; Erosion and erosion control; Global warming;
Greenhouse effect; Hydrologic cycle; Nutrient cycles; Rain forests and the
atmosphere; Soil; Soil contamination.
159
Deserts

Allaby, Michael. Deserts. New York: Facts on File, 2001.
Baylor, Byrd. The Desert Is Theirs. Reprint. New York: Aladdin Books, 1987.
Bothma, J. du P. Carnivore Ecology in Arid Lands. New York: Springer-
Verlag, 1998.
Bowers, Janice. Shrubs and Trees of the Southwest Deserts. Tucson, Ariz.:
Southwest Parks & Monuments Association, 1993.
Epple, Anne O. Field Guide to the Plants of Arizona. Helena, Mont.: Falcon,
1997.
Hare, John. The Lost Camels of Tartary: A Quest into Forbidden China. Boston:
Little, Brown, 1998.
Larson, Peggy Pickering, Lane Larson, Edward Abbey, and Iy Larson. The
Deserts of the Southwest: A Sierra Club Naturalist’s Guide. 2d ed. San Fran-
cisco: Sierra Club Books, 2000.
Raskin, Lawrence, and Debora Pearson. Fifty-two Days by Camel: My Sahara
Adventure. Toronto: Annick Press, 1998.
160
DEVELOPMENT AND ECOLOGICAL
STRATEGIES
Type of ecology: Evolutionary ecology
The relationship between ontogeny (individual development) and phylogeny (the

evolution of species and lineages) is a core concept in the study of ecology and life
sciences in general. Changes in developmental timing produce parallels between on-
togeny and phylogeny. The subject illuminates the biology of regulation, the evolu-
tion of ecological strategies, and the mechanisms for evolutionary change in form.
T he idea of a relationship between individual development, or ontog-

eny, and the evolutionary history of a race, or phylogeny, is an old one.
The concept received much attention in the nineteenth century and is often
associated with the names of Karl Ernst von Baer and Ernst Haeckel, two
prominent German biologists. It was Haeckel who coined the catchphrase
and dominant paradigm: Ontogeny recapitulates (or repeats) phylogeny.
Since Haeckel’s time, however, the relations between ontogeny and phy-
logeny have been portrayed in a variety of ways, including the reverse no-
tion that phylogeny is the succession of ontogenies. Research in the 1970’s
and 1980’s on the parallels between ontogeny and phylogeny focused on
the change of timing in developmental events as a mechanism for recapitu-
lation and on the developmental-genetic basis of evolutionary change.
Early Concepts of Biogenetic Law

During the early nineteenth century, two different concepts of parallels be-
tween development and evolution arose. The German J. F. Meckel and the
Frenchman E. R. Serres believed that a higher animal in its embryonic de-
velopment recapitulates the adult structures of animals below it on a scale
of being. Baer, on the other hand, argued that no higher animal repeats an
earlier adult stage but rather the embryo proceeds from undifferentiated
homogeneity to differentiated heterogeneity, from the general to the spe-
cific. Baer published his famous and influential four laws in 1828:
(1) The more general characters of a large group of animals appear earlier in
their embryos than the more special characters.
(2) From the most general forms, the less general are developed.
(3) Every embryo of a given animal, instead of passing through the other
forms, becomes separated from them.
(4) The embryo of a higher form never resembles any other form, only its
embryo.
161
Development and ecological strategies
By the late nineteenth century, the notion of recapitulation and Baer’s

laws of embryonic similarity were recast in evolutionary terms. Haeckel
and others established the biogenetic law: that is, ontogeny recapitulates
the adult stages of phylogeny. It was, in a sense, an updated version of the
Serres-Meckel law but differed in that the notion was valid not only for a
chain of being but also for many divergent lines of descent; ancestors had
evolved into complex forms and were now considered to be modified by
descent. More specifically, Haeckel thought of ontogeny as a short and
quick recapitulation of phylogeny caused by the physiological functions of
heredity and adaptation. During its individual development, he wrote, the
organic individual repeats the most important changes in form through
which its forefathers passed during the slow and long course of their
paleontological development. The adult stages of ancestors are repeated
during the development of descendants but crowded back into earlier
stages of ontogeny. Ontogeny is the abbreviated version of phylogeny.
These repeated stages reflect the history of the race. Haeckel considered
phylogeny to be the mechanical cause of ontogeny.
The classic example of recapitulation is the stage of development in an
unhatched bird or unborn mammal when gill slits are present. Haeckel ar-
gued that gill slits in this stage represented gill slits of the adult stage of an-
cestral fish, which in birds and mammals were pressed back into early
stages of development. This theory differed from Baer’s notion that the gill
slit in the human embryo and in the adult fish represented the same stage
in development. The gill slits, explained the recapitulationists, got from a
large adult ancestor to a small embryo in two ways: first, terminal addition
(in which stages are added to the end of an ancestral ontogeny); and sec-
ond, condensation (in which development is speeded up as ancestral fea-
tures are pushed back to earlier stages of the embryo). Haeckel also coined
another term widely used currently in another sense: “heterochrony.” He
used the term to denote a displacement in time of the appearance of one or-
gan in ontogeny before another, thus disrupting the recapitulation of phy-
logeny in ontogeny. Haeckel was not, however, interested primarily in
mechanisms or in embryology for its own sake, but rather for the informa-
tion it could provide for developing evolutionary histories.
Recapitulation in the Twentieth Century

With the rise of mechanistic experimental embryology and with the estab-
lishment of Mendelian genetics in the early twentieth century, the bio-
genetic law was largely repudiated by biologists. Descriptive embryology
was out of fashion, and the existence of genes made the two correlate laws
to recapitulation—terminal addition and condensation—untenable. One
162
of the most influential modifications for later work on the subject was
broached in a paper by Walter Garstang in 1922, in which he reformulated
the theory of recapitulation and refurbished the concept of heterochrony.
Garstang argued that phylogeny does not control ontogeny but rather
makes a record of the former: that is, phylogeny is a result of ontogeny. He
suggested that adaptive changes in a larval stage coupled with shifts in the
timing of development (heterochrony) could result in radical shifts in
adult morphology.
Stephen Jay Gould resurrected the long unpopular concept of recapitu-
lation with his book Ontogeny and Phylogeny (1977). In addition to recount-
ing the historical development of the idea of recapitulation, he made an
original contribution to defining and explicating the mechanism (het-
erochrony) involved in producing parallels between ontogeny and phy-
logeny. He argued that heterochrony—“changes in the relative time of
appearance and rate of development for characters already present in
ancestors”—was of prime evolutionary importance. He reduced Gavin de
Beer’s complex eight-mode analysis of heterochrony to two simplified
processes: acceleration and retardation. Acceleration occurs if a character
appears earlier in the ontogeny of a descendant than it did in an ancestor
because of a speeding up of development. Conversely, retardation occurs if
a character appears later in the ontogeny of a descendant than it did in an
ancestor because of a slowing down of development. To demonstrate these
concepts, Gould introduced a “clock model” in order to bring some stan-
dardization and quantification to the heterochrony concept.
He considered the primary evolutionary value of ontogeny and phylog-
eny to be in the immediate ecological advantages for slow or rapid matura-
tion rather than in the long-term changes of form. Neoteny (the opposite of
recapitulation) is the most important determinant of human evolution. Hu-
mans have evolved by retaining the young characters of their ancestors and
have therefore achieved behavioral flexibility and their characteristic form.
For example, there is a striking resemblance between some types of juvenile
apes and adult humans; this similarity for the ape soon fades in its ontogeny
as the jaw begins to protrude and the brain shrinks. Gould also insightfully
predicted that an understanding of ontogeny and phylogeny would lead
to a rapprochement between molecular and evolutionary biology.
By the 1980’s, Rudolf Raff and Thomas Kaufman found this rapproche-
ment by synthesizing embryology with genetics and evolution. Their work
focuses on the developmental-genetic mechanisms that generate evolu-
tionary change in morphology. They believe that a genetic program gov-
erns ontogeny, that the great decisions in development are made by a small
number of genes that function as switches between alternate states or path-
163
ways. When these genetic switch systems are modified, evolutionary

changes in morphology occur mechanistically. They argue further that reg-
ulatory genes—genes that control development by turning structural genes
on and off—control the timing of development, make decisions about the
fates of cells, and integrate the expression of structural genes to produce
differentiated tissue. All this plays a considerable role in evolution.
Description vs. Experimentation

Both embryology and evolution have traditionally been descriptive sci-
ences using methods of observation and comparison. By the end of the
nineteenth century, a dichotomy had arisen between the naturalistic (de-
scriptive) and the experimentalist tradition. The naturalists’ tradition
viewed the organism as a whole, and morphological studies and observa-
tions of embryological development were central to their program. Experi-
mentalists, on the other hand, focused on laboratory studies of isolated
aspects of function. A mechanistic outlook was compatible with this exper-
imental approach.
Modern embryology uses both descriptive and experimental methods.
Descriptive embryology uses topographic, histological (tissue analysis),
cytological (cell analysis), and electron microscope techniques supple-
mented by morphometric (the measurement of form) analysis. Embryos
are visualized using either plastic models of developmental stages, sche-
matic drawings, or computer simulations. Cell lineage drawings are also
used with the comparative method for phylogenies.
Experimental embryology, on the other hand, uses more invasive meth-
ods of manipulating the organism. During this field of study’s early pe-
riod, scientists subjected embryos to various changes to their normal path
of development; they were chopped into pieces, transplanted, exposed to
chemicals, and spun in centrifuges. Later, fate maps came into usage in or-
der to determine the future development of regions in the embryo. It was
found that small patches of cells on the surface of the embryo could be
stained, without damaging the cell, by applying small pieces of agar
soaked in a vital dye. One could then follow the stained cells to their even-
tual position in the gastrula. Amphibian eggs are used as material because
they are big, easy to procure, and can undergo radical experimental manip-
ulation.
Interdisciplinary Studies
Evolutionary theory primarily uses paleontology (study of the fossil rec-
ord) to study the evolutionary history of species, yet Gould also used
quantification (the clock model, for example), statistics, and ecology to un-
164
derstand the parallels between ontogeny and phylogeny. Most scientists

interested in the relationships between ontogeny and phylogeny chiefly
use comparative and theoretical methods. For example, they compare
structures in different animal groups or compare the adult structures of an
animal with the young stage of another. If similarities exist, are the lineages
similar? Are the stages in ontogenetic development similar to those of the
development of the whole species?
Yet, the study of relationships between ontogeny and phylogeny is an
interdisciplinary subject. Not only are methods from embryology and evo-
lutionary theory of help, but also, increasingly, techniques are applied
from molecular genetics. Haeckel’s method was primarily a descriptive
historical one, and he collected myriad descriptive studies of different ani-
mals. Although scientists in those days had relatively simple microscopes,
they left meticulous and detailed accounts.
A fusion of embryology, evolution, and genetics involves combining
different methods from each of the respective disciplines for the study of
the relationship between ontogeny and phylogeny. The unifying approach
has been causal-analytical, in the sense that biologists have been examin-
ing mechanisms that produce parallels between ontogeny and phylogeny
as well as the developmental-genetic basis for evolutionary change. The
methods are either technical or theoretical. The technical ones include the
use of the electron microscope, histological, cytological, and experimental
analyses; the theoretical methods include comparison, historical analysis,
observations, statistics, and computer simulation.
Ramifications Beyond Science

The relationship between ontogeny and phylogeny is one of the most im-
portant ideas in biology and a central theme in evolutionary biology. It illu-
minates the evolution of ecological strategies, large-scale evolutionary
change, and the biology of regulation. This scientific idea has also had far-
reaching influences in areas such as anthropology, political theory, litera-
ture, child development, education, and psychology.
In the late nineteenth century, embryological development was a major
part of evolutionary theory; however, that was not the case for much of the
twentieth century. Although there was some interest in embryology and
evolution from the 1920’s to the 1950’s by Garstang, J. S. Huxley, de Beer,
and Richard Goldschmidt, during the first three decades of the twentieth
century genetics and development were among the most important and
active areas in biological thought, yet there were few attempts to integrate
the two areas. It is this new synthesis of evolution, embryology, and genet-
ics that has emerged as one of the most exciting frontiers in the life sciences.
165
Although knowledge to be gained from a synthesis of development and

evolution seems not to have any immediate practical application, it can of-
fer greater insights into mechanisms of evolution, and a knowledge of evo-
lution will give similar insights into mechanisms of development. A study
of these relations and interactions also enlarges humankind’s understand-
ing of the nature of the development of individuals and their relation to the
larger historical panorama of the history of life.
Kristie Macrakis
See also: Evolution: definition and theories; Evolution: history; Extinc-

tions and evolutionary explosions; Gene flow; Genetic drift; Isolating
mechanisms; Natural selection; Nonrandom mating, genetic drift, and
mutation; Population genetics; Punctuated equilibrium vs. gradualism.

Bonner, J. T. Morphogenesis: An Essay on Development. Princeton, N.J.: Prince-
ton University Press, 1952.
De Beer, Gavin. Embryos and Ancestors. Oxford, England: Clarendon Press,
1951.
Gould, Stephen Jay. Ontogeny and Phylogeny. Cambridge, Mass.: Harvard
_______. “Ontogeny and Phylogeny—Revisited and Reunited.” BioEssays
14, no. 4 (April, 1992): 275-280.
Hall, Brian K. Evolutionary Developmental Biology. 2d ed. New York: Chap-
man & Hall, 1998.
Raff, Rudolf A., and Thomas C. Kaufman. Embryos, Genes, and Evolution:
The Developmental-Genetic Basis of Evolutionary Change. New York: Mac-
millan, 1983.
Schwartz, Jeffrey H. Sudden Origins: Fossils, Genes, and the Emergence of Spe-
cies. New York: Wiley, 1999.
166
DISPLAYS
Displays are specialized behaviors that act as communication signals within, and
occasionally between, species.
N onlinguistic forms of communication are called displays. Some dis-

plays involve, literally and simply, the visual display of a physical
feature. Among insects, for example, green or brown coloring is often used
for camouflage. Insects that are poisonous do not need camouflage and of-
ten advertise themselves with warning colors, such as black and red, or
black and orange. This is referred to as aposematic coloration or an
aposematic display.
Physical features can also indicate an individual’s sex, age, and repro-
ductive status—as do peacock tails, turkey wattles, deer antlers, the canine
teeth of male baboons, and the swollen genitals of estrous female chimpan-
zees. Many such features vary in size, shape, or color in relation to an ani-
mal’s health, hormones, or social status and are therefore referred to as sta-
tus badges or signs.
Often, meaningful physical features are highlighted by behavioral dis-
plays. A courting peacock or turkey will fan open his tail and shake it back
and forth for emphasis; a challenged buck will load plant material onto his
antlers so as to exaggerate their size; an angry baboon will curl back his lip
to expose his canine teeth; and an estrous chimpanzee will approach a
friendly male and assume a posture displaying her fertile state.
A particularly energetic or dramatic behavioral display not only calls at-
tention to a physical feature but also indicates the health and vitality of
the performer. The principle of honest signaling refers to the fact that large
and healthy individuals tend to have brighter or more contrasting colors,
make deeper-pitched and louder sounds, and produce longer, more in-
tense performances than small or weak ones. Such differences in display
quality are readily noticed by predators, potential competitors, and poten-
tial mates.
Most displays are performed by individuals and are one-way: sender to
receiver. Threat displays, however, may involve reciprocal signaling be-
tween two challengers or between two groups of challengers. Courtship
displays also may occur in groups: In some species, males gather together
to perform in what is called a lek or a lekking display. Courtship of monog-
amous species may include long sequences of frequently repeated, ritual-
167
Displays
Male peacocks fan their tails and

shake them in a courting display
to attract females. Such physical
features can also signal an
individual’s health, age, and social
status and are therefore called
“status badges.” (Digital Stock)
ized interactions in which both partners participate. Such pair-bonding

displays may continue well into the breeding season and the mateship; ini-
tially serving to familiarize the pair with one another and to synchronize
their hormones and breeding behavior, they may later serve as greeting
displays after separation.
Interpreting Displays
Charles Darwin noted that displays having opposite characteristics often
signal opposite meaning. In humans, for example, a face with upturned
corners of the mouth (a smile) signals friendliness, whereas a face with
downturned corners of the mouth (a frown) signals displeasure. In most
animals, loud, deep-pitched sounds (for example, roars and growls) indi-
cate aggression, whereas quiet, high-pitched sounds (for example, mews
and peeps) indicate anxiety or fear. Similarly, body postures exaggerating
size tend to signal dominance, whereas postures minimizing size tend
to signal submission. Darwin called his observation the principle of anti-
thesis.
Although some rules of display can be applied across species, most dis-
plays are specialized for intraspecific (within-species) communication—
male to female, parent to offspring, or dominant to subordinate—and are
therefore species-specific. That is, the ability to perform and interpret a
particular display (such as a particular birdsong) is generally characteristic
only of individuals of a particular species and is either innate (inborn) or
learned from conspecifics (individuals of the same species) during an early
critical period of development.
In order that their meaning is easily and quickly conveyed, most dis-
plays also tend to be highly ritualized; that is, they are performed only in
certain contexts and always in the same way. This consistency in commu-
168
Displays
nication prevents errors of interpretation that could be disastrous. It would

be a grave mistake, for example, to interpret an aggressive signal as a sex-
ual overture, or an alarm call (predator alert signal) as an offspring’s beg-
ging call. Mistakes of interpretation are also minimized by signal redun-
dancy; that is, messages are often conveyed simultaneously in more than
one sensory modality.
Display Modality
Displays utilize every sensory modality. Visual displays involve the use of
bright, contrasting, and sometimes changing colors; changes in body size,
shape, and posture; and what ethologists call “intention movements”—
brief, suggestive movements which reveal motivational state and likely fu-
ture actions. Auditory displays include vocal songs and calls, as well as a
variety of sounds produced by tapping, rubbing, scraping, or inflating and
deflating various parts of the body. Tactile displays include aspects of so-
cial grooming, comfort contacts (such as between littermates or parents
and offspring), and the seismic signaling of water-striders, elephants,
frogs, and spiders which vibrate, respectively, the water, ground, plants, or
web beneath them. Olfactory displays include signals from chemicals that
have been wafted into the air or water, rubbed onto objects, or deposited in
saliva, urine, or feces.
Olfaction (sense of smell) is the most primitive, and therefore the most
common and most important, sense in the animal kingdom. Species of al-
most every taxonomic group use smell to signal their whereabouts and,
generally, their sex and reproductive state. (Birds seem to be an exception.)
Animals may also use smell to identify particular individuals, to recognize
who is related to them and who is not, and to determine the relative domi-
nance status of a conspecific.
Many animals, such as this

badger, assume aggressive
postures—bared teeth, fluffed fur
to make them look larger,
growling, and a variety of other
signals—to warn off a
threatening presence such as a
predator or a human being.
(Digital Stock)
169
Displays
Chemicals used in displays are called pheromones. They may be de-

rived from waste products or hormones, acquired by ingesting certain
food items, or obtained directly from plants or other animals. Some phero-
mones not only communicate information, but also have physical effects
on their receivers.
Linda Mealey
See also: Altruism; Communication; Competition; Defense mechanisms;

Ethology; Hierarchies; Insect societies; Mammalian social systems; Migra-
tion; Mimicry; Pheromones; Poisonous animals; Predation; Reproductive
strategies; Territoriality and aggression.

Agosta, William C. Chemical Communication: The Language of Pheromones.
New York: Scientific American Library, 1992.
Bailey, Winston. Acoustic Behaviour in Insects. New York: Chapman and
Hall, 1991.
Eibl-Eibesfeldt, I. Ethology: The Biology of Behavior. Translated by Erich
Klinghammer. 2d ed. New York: Holt, Reinhart and Winston, 1975.
Guthrie, R. Dale. Body Hot Spots: The Anatomy of Human Social Organs and
Behavior. New York: Van Nostrand Reinhold, 1976.
Johnsgard, Paul A. Arena Birds: Sexual Selection and Behavior. Washington,
D.C.: Smithsonian Institution Press, 1994.
Morris, Desmond. Animalwatching. New York: Crown, 1990.
Owen, Denis. Camouflage and Mimicry. Chicago: University of Chicago
Press, 1980.
170
ECOLOGY: DEFINITION
Type of ecology: Theoretical ecology
Ecology is the study of the relationships of organisms to their environments. By ex-

amining those relationships in natural ecosystems, ecologists can discover princi-
ples that help humankind understand its own role on the planet.
E cology is the study of how organisms relate to their natural environ-

ments. The two principal concerns of ecologists are the distribution
and abundance of organisms: Why are animals, plants, and other organ-
isms found where they are, and why are some common and others rare?
These questions have their roots in the theory of evolution. In fact, it is dif-
ficult (and not often worthwhile) to separate modern ecological matters
from the concerns of evolutionary biologists. Ecology can be divided ac-
cording to several levels of organization: the individual organism, the pop-
ulation, the community, and the ecosystem.
Individual Organisms
An ecologist views organisms as consequences of past natural selection
brought about by their environments. That is, each organism represents an
array of adaptations that can provide insight into the environmental pres-
sures that resulted in its present form. Adaptations of organisms are also
revealed by other features, such as the range of temperature an organism
can tolerate, the amount of moisture it requires, or the variety of food it can
exploit. Food and space for living are considered resources; factors such as
temperature, light, and moisture are conditions that determine the rate of
resource utilization. When ecologists have discovered the full range of re-
sources and conditions necessary for an organism’s existence, they have
discovered its niche.
Many species, such as many insects and plants, have a large reproduc-
tive output. This compensates for high mortality imposed by natural selec-
tion. Other species, such as large mammals and birds, have fewer off-
spring. Many of these animals care for their young, thus increasing the
chances that their offspring will survive to reproduce. These are two differ-
ent strategies for success, based upon the principle that organisms have a
finite energy budget. Energy acquired from food (animals) or sunlight
(plants) must be partitioned among growth, maintenance, and reproduc-
tion. The greater the energy allocated to the care of offspring, for example,
the fewer the offspring that can be produced.
171
Ecology: definition
The concept of an energy budget is a key to understanding evolution-

ary strategies of organisms, as well as the energetics of ecosystems. The
amount of energy fixed and stored by an organism is called net production;
this is the energy used for growth and reproduction. Net production is the
difference between gross production (the amount of energy assimilated)
and respiration (metabolic maintenance cost). The greater the respiration,
the less energy will be left over for growth and reproduction. Endothermic
animals, which physiologically regulate their body heat (mammals and
birds), have a very high respiration rate relative to ectotherms (reptiles,
amphibians, fish, and invertebrates), which cannot. Among endotherms,
smaller animals have higher respiration rates than larger ones, because the
ratio of body surface area (the area over which heat is exchanged with the
environment) to volume (the size of the “furnace”) decreases with increas-
ing body size.
Populations
Although single organisms can be studied with regard to adaptations, in
nature most organisms exist in populations rather than as individuals.
Some organisms reproduce asexually (that is, by forming clones), so that a
single individual may spawn an entire population of genetically identical
individuals. Populations of sexually reproducing organisms, however,
have the property of genetic variability, since not all individuals are iden-
tical. That is, members of a population have slightly different niches and
will therefore not all be equally capable of living in a given environment.
This is the property upon which Charles Darwin’s theory of natural selec-
tion depends: Because not all individuals are identical, some will have
greater fitness than others. Those with superior fitness will reproduce in
greater numbers and therefore will contribute more genes to successive
generations. In nature, many species consist of populations occupying
more than a single habitat. This constitutes a buffer against extinction: If
one habitat is destroyed, the species will not go extinct, because it exists in
other habitats.
Two dynamic features of populations are growth and regulation. Growth
is simply the difference between birth and death rates, which can be posi-
tive (growing), negative (declining), or zero (in equilibrium). Every species
has a genetic capacity for exponential (continuously accelerating) increase,
which will express itself to varying degrees depending on environmental
conditions: A population in its ideal environment will express this capacity
more nearly than one in a less favorable environment. The rate of growth
of a population is affected by its age structure—the proportion of individu-
als of different ages. For example, a population that is growing rapidly will
172
Ecology: definition
have a higher proportion of juvenile individuals than one that is growing

more slowly.
Populations may be regulated (so that they have equal birth and death
rates) by a number of factors, all of which are sensitive to changes in popu-
lation size. A population may be regulated by competition among its mem-
bers for the resource that is in shortest supply (limiting). The largest popu-
lation that can be sustained by the available resources is called the carrying
capacity of the environment. A population of rodents, for example, might be
limited by its food supply such that as the population grows and food runs
out, the reproductive rate declines. Thus, the effect of food on population
growth depends upon the population size relative to the limiting resource.
Similarly, parasites that cause disease spread faster in large, dense popula-
tions than in smaller, more diffuse ones. Predators can also regulate popula-
tions of their prey by responding to changes in prey availability. Climate and
catastrophic events such as storms may severely affect populations, but their
effect is not dependent upon density and is thus not considered regulatory.
Communities
Communities of organisms are composed of many populations that may
interact with one another in a variety of ways: predation, competition, mu-
tualism, parasitism, and so on. The composition of communities changes
over time through the process of succession. In terrestrial communities,
bare rock may be weathered and broken down by bacteria and other or-
ganisms until it becomes soil. Plants can then invade and colonize this
newly formed soil, which in turn provides food and habitat for animals.
The developing community goes through a series of stages, the nature of
which depends on local climatic conditions, until it reaches a kind of equi-
librium. In many cases this equilibrium stage, called climax, is a mature
forest. Aquatic succession essentially is a process of becoming a terrestrial
community. The basin of a lake, for example, will gradually be filled with
silt from terrestrial runoff and accumulated dead organic material from
populations of organisms within the lake itself.
Competition occurs between, as well as within, species. Two species are
said to be in competition with each other if and only if they share a resource
that is in short supply. If, however, they merely share a resource that is
plentiful, then they are not really competing for it. Competition is thought
to be a major force in determining how many species can coexist in natural
communities. There are a number of alternative hypotheses, however,
which involve such factors as evolutionary time, productivity (the energy
base for a community), heterogeneity of the habitat, and physical harsh-
ness of the environment.
173
Ecology: definition
Predator-prey interactions are those in which the predator benefits from

killing and consuming its prey. These differ from most parasite-host inter-
actions in that parasites usually do not kill their hosts (a form of suicide for
creatures that live inside other creatures). Similarly, most plant-eating ani-
mals (herbivores) do not kill the plants on which they feed. Many ecolo-
gists classify herbivores as parasites for this reason. There are exceptions,
such as birds and rodents that eat seeds, and these can be classified as legit-
imate predator-prey interactions. Predators can influence the number of
species in a community by affecting competition among their prey: If pop-
ulations of competing species are lowered by predators so that they are be-
low their carrying capacities, then there may be enough resources to sup-
port colonization by new species.
In many cases, the interaction between two species is mutually benefi-
cial. Mutualism is often thought to arise as a result of closely linked evolu-
tionary histories (coevolution) of different species. Termites harbor proto-
zoans in their guts that produce an enzyme that breaks down cellulose in
wood. The protozoans thus are provided with a habitat, and the termites in
turn are able to derive nourishment from wood. Some acacia trees in the
tropics have hollow thorns which provide a habitat for ants. In return, the
ants defend the trees from other insects which would otherwise damage or
defoliate them.
Ecosystems
Ecosystems consist of several trophic levels, or levels at which energy is ac-
quired: primary producers, consumers, and decomposers. Primary pro-
ducers are green plants that capture solar energy and transform it, through
the process of photosynthesis, into chemical energy. Organisms that eat
plants (herbivores) or animals (carnivores) to obtain their energy are col-
lectively called consumers. Decomposers are those consumers, such as
bacteria and fungi, that obtain energy by breaking down dead bodies of
plants and animals. These trophic levels are linked together into a struc-
ture called a food web, in which energy is transferred from primary pro-
ducers to consumers and decomposers, until finally all is lost as heat. Each
transfer of energy entails a loss (as heat) of at least 90 percent, which means
that the total amount of energy available to carnivores in an ecosystem is
substantially less than that available to herbivores.
As with individual organisms, ecosystems and their trophic levels have
energy budgets. The net production of one trophic level is available to the
next-higher trophic level as biomass (mass of biological material). Plants
have higher net productivity (rates of production) than animals because
their metabolic maintenance cost is lower relative to gross productivity;
174
Ecology: definition
herbivores often have higher net productivity than predators for the same
reason. For the community as a whole, net productivity is highest during
early successional stages, since biomass is being added more rapidly than
later on, when the community is closer to climax equilibrium.
In contrast to the unidirectional flow of energy, materials are conserved
and recycled from dead organisms by decomposers to support productiv-
ity at higher trophic levels. Carbon, water, and mineral nutrients required
for plant growth are cycled through various organisms within an ecosys-
tem. Materials and energy are also exchanged among ecosystems: There
is no such thing in nature as a “closed” ecosystem that is entirely self-
contained.
Methods of Studying Ecology

The science of ecology is necessarily more broadly based than most biolog-
ical disciplines; consequently, there is more than one approach to it. Eco-
logical studies fall into three categories: descriptive, experimental, and
mathematical.
Descriptive ecology is concerned with describing natural history, usu-
ally in qualitative terms. The study of adaptations, for example, is descrip-
tive in that one can measure the present “value” of an adaptive feature, but
one can only conjecture as to the history of natural selection that was re-
sponsible for it. On the other hand, there are some patterns discernible in
nature for which hypotheses can be constructed and tested by statistical in-
ference. For example, the spatial distribution (dispersion) of birds on an is-
land may be random, indicating no biological interaction among them. If
the birds are more evenly spaced (uniform dispersion) than predicted as-
suming randomness, however, then it might be inferred that the birds are
competing for space; they are exhibiting territorial exclusion of one an-
other. Such “natural experiments,” as they are called, depend heavily upon
the careful design of statistical tests.
Experimental ecology is no different from any other experimental disci-
pline; hypotheses are constructed from observations of nature, controlled
experiments are designed to test them, and conclusions are drawn from the
results of the experiments. The basic laboratory for an ecologist is the field.
Experiments in the field are difficult because it is hard to isolate and ma-
nipulate variable factors one at a time, which is a requisite for any good ex-
periment in science. A common experiment that is performed to test for re-
source limitation in an organism is enhancement of that resource. If food,
for example, is thought to be in short supply (implying competition), one
section of the habitat is provided more food than is already present; an-
other section is left alone as a control. If survivorship, growth, or repro-
175
Ecology: definition
ductive output is higher in the enhanced portion of the habitat than in the
control area, the researcher may infer that the organisms therein were
food-limited. Alternatively, an ecologist might have decreased the density
of organisms in one portion of the habitat, which might seem equivalent to
increasing food supply for the remaining organisms, except that it repre-
sents a change in population density as well. Therefore, this second design
will not allow the researcher to differentiate between the possibly separate
effects of food level and simple population density on organisms in the
habitat.
Mathematical ecology relies heavily upon computers to generate mod-
els of nature. A model is simply a formalized, quantitative set of hypothe-
ses constructed from sets of assumptions of how things happen in nature.
A model of population growth might contain assumptions about the age
structure of a population, its genetic capacity for increase, and the average
rate of resource utilization by its members. By changing these assump-
tions, scientists can cause the model population to behave in different
ways over time. The utility of such modeling is limited to the accuracy of
the assumptions employed.
Modern ecology is concerned with integrating these different approaches,
all of which have in common the goal of predicting the way nature will be-
have in the future, based upon how it behaves in the present. Description
of natural history leads to hypotheses that can be tested experimentally,
which in turn may allow the construction of realistic mathematical (quanti-
tative) models of how nature works.
Human Impacts and Applications

People historically have viewed nature as an adversary. The “conquest of
nature” has traditionally meant human encroachment on natural ecosys-
tems, usually without benefit of predictive knowledge. Such environmen-
tal problems as pollution, species extinction, and overpopulation can be
viewed as experiments performed on a grand scale without appropriate
controls. The problem with such experiments is that the outcomes might
be irreversible. A major lesson of ecology is that humans are not separate
from nature; we are constrained by the same principles as are other organ-
isms on the earth. One object of ecology, then, is to learn these principles so
that they can be applied to our portion of the earth’s ecosystem.
Populations that are not regulated by predators, disease, or food limita-
tion grow exponentially. The human population, on a global scale, has
grown this way. All the wars and famines in history have scarcely made a
dent in this growth pattern. Humankind has yet to identify its carrying ca-
pacity on a global scale, although regional famines certainly have provided
176
Ecology: definition
insights into what happens when local carrying capacity is exceeded. The
human carrying capacity needs to be defined in realistic ecological terms,
and such constraints as energy, food, and space must be incorporated into
the calculations. For example, knowledge of energy flow teaches that there
is more energy at the bottom of a food web (producers) than at successively
higher trophic levels (consumers), which means that more people could be
supported as herbivores than as carnivores.
The study of disease transmission, epidemiology, relies heavily on
ecological principles. Population density, rates of migration among epi-
demic centers, physiological tolerance of the host, and rates of evolution
of disease-causing parasites are all the subjects of ecological study.
An obvious application of ecological principles is conservation. Before
habitats for endangered species can be set aside, for example, their ecologi-
cal requirements, such as migratory routes, breeding, and feeding habits,
must be known. This also applies to the introduction (intentional or acci-
dental) of exotic species into habitats. History is filled with examples of in-
troduced species that caused the extinction of native species. Application
of ecological knowledge in a timely fashion, therefore, might prevent spe-
cies from becoming endangered in the first place.
One of the greatest challenges humans face is the loss of habitats world-
wide. This is especially true of the tropics, which contain most of the
earth’s species of plants and animals. Species in the tropics have narrow
niches, which means that they are more restricted in range and less tolerant
of change than are many temperate species. Therefore, destruction of tropi-
cal habitats, such as rain forests, leads to rapid species extinction. These
species are the potential sources of many pharmaceutically valuable drugs;
further, they are a genetic record of millions of years of evolutionary his-
tory. Tropical rain forests also are prime sources of oxygen and act as a
buffer against carbon dioxide accumulation in the atmosphere. Ecological
knowledge of global carbon cycles permits the prediction that destruction
of these forests will have a profound impact on the quality of the air.
Lawrence E. Hurd
See also: Balance of nature; Biomes: determinants; Biomes: types; Bio-

sphere concept; Deep ecology; Ecosystems: definition and history; Sustain-
able development.

uals, Populations, and Communities. 3d ed. Cambridge, Mass.: Blackwell
Science, 1996.
177
Ecology: definition
Bush, M. B. Ecology of a Changing Planet. 2d ed. Upper Saddle River, N.J.:

Prentice-Hall, 2000.
Carson, Rachel. Silent Spring. Boston: Houghton Mifflin, 1962.
Elton, Charles. Animal Ecology. New York: Macmillan, 1927.
Hutchinson, G. Evelyn. The Ecological Theater and the Evolutionary Play.
New Haven, Conn.: Yale University Press, 1969.
Krebs, Charles J. Ecological Methodology. 2d ed. Menlo Park, Calif.: Benjamin/
Cummings, 1999.
_______. Ecology: The Experimental Analysis of Distribution and Abundance.
5th ed. San Francisco: Benjamin/Cummings, 2001.
_______. The Message of Ecology. New York: Harper & Row, 1987.
Pianka, Eric R. Evolutionary Ecology. 6th ed. San Francisco: Benjamin/Cum-
mings, 2000.
Ricklefs, Robert E. Ecology. 4th ed. New York: Chiron Press, 1999.
178
ECOLOGY: HISTORY
Type of ecology: History of ecology
As a formal discipline, ecology, the science that studies the relationships among or-
ganisms and their biotic and abiotic environments, is a relatively new science,
which became a focus of study at about the same time evolutionary theories were
being proposed.
T he study of ecological topics arose in ancient Greece, but these studies

were part of a catch-all science called natural history. The earliest at-
tempt to organize an ecological science separate from natural history was
made by Carolus Linnaeus in his essay Oeconomia Naturae (1749; The Econ-
omy of Nature), which focused on the balance of nature and the environ-
ments in which various natural communities existed. Although the essay
was well known, the eighteenth century was dominated by biological ex-
ploration of the world, and the science of ecology did not develop.
Early Ecological Studies

The study of fossils led some naturalists to conclude that many species
known only as fossils must have become extinct. However, Jean-Baptiste
Lamarck argued in his Philosophie zoologique (1809; Zoological Philosophy,
1914) that fossils represented the early stages of species that evolved into
different species that were still living. In order to refute this claim, geolo-
gist Charles Lyell mastered the science of biogeography and used it to ar-
gue that species do become extinct and that competition from other species
seemed to be the main cause. English naturalist Charles Darwin’s book On
the Origin of Species by Means of Natural Selection (1859) blends his own re-
searches with the influence of Linnaeus and Lyell in order to argue that
some species do become extinct, but existing species have evolved from
earlier ones. Lamarck had underrated and Lyell had overrated the impor-
tance of competition in nature.
Although Darwin’s book was an important step toward ecological sci-
ence, he and his colleagues mainly studied evolution rather than ecology.
German evolutionist Ernst Haeckel realized the need for an ecological sci-
ence and coined the name oecologie in 1866. It was not until the 1890’s, how-
ever, that steps were taken to organize this science. Virtually all of the early
ecologists were specialists in the study of particular groups of organisms,
and it was only in the late 1930’s that some efforts were made to write text-
books covering all aspects of ecology. Since the 1890’s, many ecologists
179
Ecology: history
have viewed themselves as plant ecologists, animal ecologists, marine bi-

ologists, or limnologists. Limnology is the study of freshwater aquatic en-
vironments.
Nevertheless, general ecological societies were established. The first
was the British Ecological Society, which was founded in 1913 and began
publishing the Journal of Ecology in the same year. Two years later, ecolo-
gists in the United States and Canada founded the Ecological Society of
America, which began publishing Ecology as a quarterly journal in 1920; in
1965 Ecology began appearing bimonthly. Other national societies have
since been established. More specialized societies and journals also began
appearing. For example, the Limnological Society of America was estab-
lished in 1936 and expanded in 1948 into the American Society of Limnol-
ogy and Oceanography. It publishes the journal Limnology and Oceanogra-
phy. These and now many other professional organizations, both academic
and applied, sponsor not only journals but also Web sites, reports, and
symposia.
Although Great Britain and Western Europe were active in establishing
the study of ecological sciences, it was difficult for their trained ecologists
to obtain full-time employment that utilized their expertise. European uni-
versities were mostly venerable institutions with fixed budgets; they al-
ready had as many faculty positions as they could afford, and these were
all allocated to the older arts and sciences. Governments employed few, if
any, ecologists. The situation was more favorable in the United States, Can-
ada, and Australia, where universities were still growing. In the United
States, the universities that became important for ecological research and
the training of new ecologists were mostly in the Midwest. The reason was
that eastern universities were similar to European ones in being well estab-
lished, with scientists in traditional fields.
Ecology After 1950

Ecological research in the United States was not well funded until after
World War II. With the advent of the Cold War, science was suddenly con-
sidered important for national welfare. In 1950 the U.S. Congress estab-
lished the National Science Foundation, and ecologists were able to make
the case for their research along with that of the other sciences. The Atomic
Energy Commission had already begun to fund ecological research by
1947, and under its patronage the Oak Ridge National Laboratory and the
University of Georgia gradually became important centers for radiation
ecology research.
Another important source of research funds was the International Bio-
logical Program (IBP), which, though international in scope, depended
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Ecology: history
upon national research funds. It got under way in the United States in 1968
and was still producing publications in the 1980’s. Even though no new
funding sources were created for the IBP, its existence meant that more re-
search money flowed to ecologists than otherwise would have.
Ecologists learned to think big. Computers became available for ecolog-
ical research shortly before the IBP got under way, and so computers and
the IBP became linked in ecologists’ imaginations. Earth Day, established
in 1970, helped awaken Americans to the environmental crisis. The IBP en-
couraged a variety of studies, but in the United States, studies of biomes
(large-scale environments) and ecosystems were most prominent. The bio-
me studies were grouped under the headings of desert, eastern deciduous
forest, western coniferous forest, grassland, and tundra (a proposed tropi-
cal forest program was never funded). When the IBP ended, a number of
its biome studies continued at a reduced level.
Ecosystem studies are also large-scale, at least in comparison with
many previous ecological studies, though smaller in size than a biome. The
goal of ecosystem studies was to gain a total understanding of how an eco-
system—such as a lake, river valley, or forest—works. IBP funds enabled
students to collect data, which computers processed. However, ecologists
could not agree on what data to collect, how to compute outcomes, and
how to interpret the results. Therefore, thinking big did not always pro-
duce impressive results.
Plant Ecology
Because ecology is enormous in scope, it was bound to have growing
pains. It arose at the same time as the science of genetics, but because ge-
netics is a cohesive science, it reached maturity much sooner than ecology.
Ecology can be subdivided in a wide variety of ways, and any collection of
ecology textbooks will show how diversely it is organized by different
ecologists. Nevertheless, self-identified professional subgroups tend to
produce their own coherent findings.
Plant ecology progressed more rapidly than other subgroups and has
retained its prominence. In the early nineteenth century, German naturalist
Alexander von Humboldt’s many publications on plant geography in rela-
tion to climate and topography were a powerful stimulus to other bota-
nists. By the early twentieth century, however, the idea of plant communi-
ties was the main focus for plant ecologists. Henry Chandler Cowles began
his studies at the University of Chicago in geology but switched to botany
and studied plant communities on the Indiana dunes of Lake Michigan.
He received his doctorate in 1898 and stayed at that university as a plant
ecologist. He trained others in the study of community succession.
181
Ecology: history
Frederic Edward Clements received his doctorate in botany in 1898

from the University of Nebraska. He carried the concept of plant com-
munity succession to an extreme by taking literally the analogy between
the growth and maturation of an organism and that of a plant com-
munity. His numerous studies were funded by the Carnegie Institute in
Washington, D.C., and even ecologists who disagreed with his theoreti-
cal extremes found his data useful. Henry Allan Gleason was skeptical;
his studies indicated that plant species that have similar environmental
needs compete with one another and do not form cohesive communities.
Although Gleason first expressed his views in 1917, Clements and his
disciples held the day until 1947, when Gleason’s individualistic concept
received the support of three leading ecologists. Debates over plant succes-
sion and the reality of communities helped increase the sophistication of
plant ecologists and prepared them for later studies on biomes, ecosys-
tems, and the degradation of vegetation by pollution, logging, and agricul-
ture.
Marine Ecology
Marine ecology is viewed as a branch of either ecology or oceanography.
Early studies were made either from the ocean shore or close to shore be-
cause of the great expense of committing oceangoing vessels to research.
The first important research institute was the Statione Zoologica at Naples,
Italy, founded in 1874. Its successes soon inspired the founding of others in
Europe, the United States, and other countries. Karl Möbius, a German zo-
ologist who studied oyster beds, was an important pioneer of the commu-
nity concept in ecology. Great Britain dominated the seas during the nine-
teenth century and made the first substantial commitment to deep-sea
research by equipping the HMS Challenger as an oceangoing laboratory
that sailed the world’s seas from 1872 to 1876. Its scientists collected so
many specimens and so much data that they called upon marine scientists
in other countries to help them write the fifty large volumes of reports
(1885-1895). The development of new technologies and the funding of new
institutions and ships in the nineteenth century enabled marine ecologists
to monitor the world’s marine fisheries and other resources and provide
advice on harvesting marine species.
Limnology is the scientific study of bodies of fresh water. The Swiss
zoologist François A. Forel coined the term and also published the first
textbook on the subject in 1901. He taught zoology at the Académie de
Lausanne and devoted his life’s researches to understanding Lake Ge-
neva’s characteristics and its plants and animals. In the United States in the
early twentieth century, the University of Wisconsin became the leading
182
Ecology: history
center for limnological research and the training of limnologists, and it has
retained that preeminence. Limnology is important for managing freshwa-
ter fisheries and water quality.
Frank N. Egerton
See also: Ecology: history; Ecosystems: definition and history; Evolution:

history.

Allen, Timothy F. H., and Thomas W. Hoekstra. Toward a Unified Ecology.
New York: Columbia University Press, 1992.
Hynes, H. Patricia. The Recurring Silent Spring. New York: Pergamon Press,
1989.
Leuzzi, Linda. Life Connections: Pioneers in Ecology. Danbury, Conn.: Frank-
lin Watts, 2000.
183
ECOSYSTEMS:
DEFINITION AND HISTORY
Types of ecology: Ecosystem ecology; History of ecology; Theoretical

ecology
The ecosystem is the fundamental concept in ecology: the basic unit of nature, con-
sisting of the complex of interacting organisms inhabiting a region with all the
nonliving physical factors that make up their environment. Ecologists study
structural and functional relationships of ecosystem components to be able to pre-
dict how the system will respond to natural change and human disturbance.
T he ecosystem is essentially an abstract organizing unit superimposed

on the landscape to help ecologists study the form and function of the
natural world. An ecosystem consists of one or more communities of inter-
acting organisms and their physical environment. Ecosystems have no dis-
tinct boundaries; thus the size of any particular ecosystem should be in-
ferred from the context of the discussion. Individual lakes, streams, or
strands of trees can be described as distinct ecosystems, as can the entire
North American Great Lakes region. Size and boundaries are arbitrary be-
cause no ecosystem stands in complete isolation from those that surround
it. A lake ecosystem, for example, is greatly affected by the streams that
flow into it and by the soils and vegetation through which these streams
flow. Energy, organisms, and materials routinely migrate across whatever
perimeters the ecologist may define. Thus, investigators are allowed con-
siderable latitude in establishing the scale of the ecosystem they are study-
ing. Whatever the scale, though, the importance of the ecosystem concept
is that it forces ecologists to treat organisms not as isolated individuals or
species but in the context of the structural and functional conditions of
their environment.
Development of the Ecosystem Concept

Antecedents to the ecosystem concept may be traced back to one of Amer-
ica’s first ecologists, Stephen Alfred Forbes, an eminent Illinois naturalist
who studied food relationships among birds, fish, and insects. During the
course of his investigations, Forbes recognized in 1880 that full knowledge
of organisms and their response to disturbances would come only from
more concentrated research on their interactions with other organisms and
with their inorganic (nonliving) physical surroundings. In 1887, Forbes
184
Ecosystems: definition and history
suggested that a lake could be viewed as a discrete system for study: a mi-
crocosm. A lake could serve as a scale model of nature that would help bi-
ologists understand more general functional relationships among organ-
isms and their environment. Forbes explained how the food supply of a
single species, the largemouth bass, was dependent either directly or indi-
rectly upon nearly all the fauna and much of the flora of the lake. There-
fore, whenever even one species was subjected to disturbance from outside
the microcosm, the effects would probably be felt throughout the commu-
nity.
In 1927, British ecologist Charles Elton incorporated ideas introduced
by Forbes and other fishery biologists into the twin concepts of the food
chain and the food web. Elton defined a food chain as a series of linkages
connecting basic plants, or food producers, to herbivores and their various
carnivorous predators, or consumers. Elton used the term “food cycle” in-
stead of “food web,” but his diagrams reveal that his notion of a food cycle—
that it is simply a network of interconnecting food chains—is consistent
with the modern term.
Elton’s diagrams, which traced various pathways of nitrogen through
the community, paved the way for understanding the importance of the
cycling of inorganic nutrients such as carbon, nitrogen, and phosphorus
through ecosystems, a process that is known as biogeochemical cycling.
Very simply, Elton illustrated how bacteria could make nitrogen available
to algae at the base of the food chain. The nitrogen then could be incorpo-
rated into a succession of ever larger consumers until it reached the top of
the chain. When the top predators died, decomposer organisms would re-
turn the nitrogen to forms that could eventually be taken up again by
plants and algae at the base of the food chain, thus completing the cycle.
Elton’s other key contribution to the ecosystem concept was his articu-
lation of the pyramid of numbers, the idea that small animals in any given
community are far more common than large animals. Organisms at the
base of a food chain are numerous, and those at the top are relatively
scarce. Each level of the pyramid supplies food for the level immediately
above it—a level consisting of various species of predators that generally
are larger in size and fewer in number. That level, in turn, serves as prey for
a level of larger, more powerful predators, fewer still in number. A graph of
this concept results in a pyramidal shape of discrete levels, which today are
called trophic (feeding) levels.
Ecosystems in Twentieth Century Thought

Although the basic concept of an ecosystem had been recognized by
Forbes as early as 1880, it was not until 1935 that British ecologist Arthur G.
185
Tansley coined the term. Though he acknowledged that ecologists were

primarily interested in organisms, Tansley declared that organisms could
not be separated from their physical environments, as organism and envi-
ronment formed one complete system. As Forbes had pointed out half a
century earlier, organisms were inseparably linked to their nonliving envi-
ronment. Consequently, ecosystems came to be viewed as consisting of
two fundamental parts: the biotic, or living components, and the abiotic, or
nonliving components.
No one articulated this better than Raymond Lindeman, a limnologist
(freshwater biologist) from Minnesota who in 1941 skillfully integrated the
ideas of earlier ecological scientists when he published an elegant ecosys-
tem study of Cedar Bog Lake. Lindeman’s classic work set the stage for de-
cades of research that centered on the ecosystem as the primary organizing
unit of study in ecology. Drawing on the work of his mentor, G. Evelyn
Hutchinson, and Charles Elton, Arthur Tansley, and other scientists, Lin-
deman explained how ecological pyramids were a necessary result of en-
ergy transfers from one trophic level to the next.
By analyzing ecosystems in this manner, Lindeman was able to answer
a fundamental ecological question that had been posed fourteen years ear-
lier by Elton: Why were the largest and most powerful animals, such as po-
lar bears, sharks, and tigers, so rare? Elton had thought the relative scarcity
of top predators was due to their lower rates of reproduction. Lindeman
corrected this misconception by explaining that higher trophic levels held
fewer animals not because of their reproductive rates but because of a loss
of chemical energy with each step up the pyramid. It could be looked upon
as a necessary condition of the second law of thermodynamics: Energy
transfers yield a loss or degradation of energy. The predators of one food
level could never completely extract all the energy from the level below.
Some energy would always be lost to the environment through respira-
tion, some energy would not be assimilated by the predators, and some en-
ergy simply would be lost to decomposer chains when potential prey died
of nonpredatory causes. This meant that each successive trophic level had
substantially less chemical energy available to it than was transferred to
the one below and, therefore, could not support as many animals.
Ecologists soon expanded the principle of Elton’s pyramid of numbers
to model other ecosystem processes. They found, for example, that the
flow of chemical energy through an ecosystem could be characterized as
an energy pyramid; the biomass (the weight of organic material, as in plant
or animal tissue) in a community could be plotted in a pyramid of biomass.
Collectively, such pyramidal models became known as ecological pyra-
mids.
186
Energy Production and Transmission

Lindeman subdivided the biotic components of ecosystems into produc-
ers, consumers, and decomposers. Producers (also known as autotrophs)
produce their own food from compounds in their environment. Green
plants are the main producers in terrestrial ecosystems; algae are the most
common producers in aquatic ecosystems. Both plants and algae are pro-
ducers that use sunlight as energy to make food from carbon dioxide
and water in the process of photosynthesis. During photosynthesis, plants,
algae, and certain bacteria capture the sun’s energy in chlorophyll mol-
ecules. (Chlorophyll is a pigment that gives plants their green color.) This
energy, in turn, is used to synthesize energy-rich compounds such as
glucose, which can be used to power activities such as growth, main-
tenance, and reproduction or can be stored as biomass for later use. These
energy-rich compounds can also be passed on in the form of biomass
from one organism to another, as when animals (primary consumers)
graze on plants or when decomposers break down detritus (dead organic
matter).
The energy collected by green plants is called primary production be-
cause it forms the first level at the base of the ecological pyramid. Total
photosynthesis is represented as gross primary production. This is the
amount of the sun’s energy actually assimilated by autotrophs. The rate
of this production of organic tissue by photosynthesis is called primary
productivity. Plants, however, need to utilize some of the energy they pro-
duce for their own growth, maintenance, and reproduction. This energy
becomes available for such activities through respiration, which essen-
tially is a chemical reversal of the process of photosynthesis. As a result,
not all the energy assimilated by autotrophs is available to the consumers
in the next trophic level of the pyramid. Consequently, respiration costs
generally are subtracted from gross primary production to determine the
net primary production, the chemical energy actually available to primary
consumers.
Measuring Ecosystem Productivity

The carrying capacity for all the species supported by an ecosystem ulti-
mately depends upon the system’s net primary productivity. By knowing
the productivity, ecologists can, for example, estimate the number of herbi-
vores that an ecosystem can support. Consequently ecosystem ecologists
have developed a variety of methods to measure the net primary produc-
tivity of different systems. Productivity is generally expressed in kilo-
calories per square meter per year when quantifying energy, and in grams
per square meter per year when quantifying biomass.
187
Production in aquatic ecosystems may be measured by using the light

and dark bottle method. In this technique two bottles containing samples
of water and the natural phytoplankton population are suspended for
twenty-four hours at a given depth in a body of water. One bottle is dark,
permitting respiration but no photosynthesis by the phytoplankton. The
other is clear and therefore permits both photosynthesis and respiration.
The light bottle provides a measure of net production (photosynthesis mi-
nus respiration) if the quantity of oxygen is measured before and after the
twenty-four-hour period. (The amount of oxygen produced by photosyn-
thesis is proportional to the amount of organic matter fixed.) Measuring
the amount of oxygen in the dark bottle before and after the run provides
an estimate of respiration, since no photosynthesis can occur in the dark.
Combining net production from the light bottle with total respiration from
the dark bottle yields an estimate of gross primary production.
Other studies have concentrated on quantifying the rate of movement
of energy and materials through ecosystems. Investigations begun in the
1940’s and 1950’s by the Atomic Energy Commission to track radioactive
fallout were eventually diverted into studies of ecosystems that demon-
strated how radionuclides moved through natural environments by
means of food-chain transfers. This research confirmed the interlocking
nature of all organisms linked by the food relationship and eventually
yielded rates at which both organic and inorganic materials could be cy-
cled through ecosystems. As a result of such studies, the Radiation Ecology
Section at Oak Ridge National Laboratory in Tennessee became estab-
lished as a principal center for systems ecology.
Ecologists sometimes extend the temporal boundaries of their studies
by utilizing the methods of paleoecology (the use of fossils to study the na-
ture of ecosystems in the past). Research of this type generally centers on
the analysis of lake sediments, whose layers often hold centuries of ecosys-
tem history embodied in the character and abundance of pollen grains, di-
atoms, fragments of zooplankton, and other organic microfossils.
More general trends in the methods of studying ecosystems include a
continuing emphasis on quantitative methods, often using increasingly so-
phisticated computer modeling techniques to simulate ecosystem func-
tions. Equally significant is a trend toward a “big science” approach, mod-
eled on the Manhattan Project, which employed teams of investigators
working on different problems related to nuclear fission in different parts
of the country. The well-known international biological program, the Hub-
bard Brook project in New Hampshire, and continuing projects on long-
term ecological research all serve as examples of ecosystem studies that in-
volve teams of researchers from a wide range of disciplines.
188
Responding to Disturbance
One of the practical benefits of studying ecosystems derives from natural-
ist Stephen Forbes’s suggestion, made in 1880, that the knowledge from bi-
ological research be used to predict the response of organisms to distur-
bance. When disturbance is caused by natural events such as droughts,
floods, or fires, ecologists can use their knowledge of the structure and
function of ecosystems to help resource managers plan for the subsequent
recolonization and succession of species.
The broad perspective of the ecosystem approach becomes particularly
useful in examining the effects of certain toxic compounds because of the
complexity of their interaction within the environment. The synergistic ef-
fects that sometimes occur with toxic substances can produce pronounced
impacts on ecosystems already stressed by other disturbances. For example,
after the atmosphere deposits mercury on the surface of a lake, the pollutant
eventually settles in the sediments where bacteria make it available to or-
ganisms at the base of the food chain. The contaminant then bioaccumulates
as it is passed on to organisms such as fish and fish-eating birds at higher
trophic levels. Synergistic effects occur in lakes already affected by acid de-
position; researchers have found that acidity somehow stimulates microbes
to increase the bioavailability of the mercury. Thus, aquatic ecosystems
that have become acidified through atmospheric processes may stress their
flora and fauna even further by enhancing the availability of mercury from
atmospheric fallout. The complexity of such interactions demands re-
search at the ecosystem level, and ecosystem studies are prerequisite for
prudent public policy actions on environmental contaminants.
Robert Lovely
See also: Biomes: determinants; Biomes: types; Communities: ecosystem

interactions; Ecology: history; Ecosystems: studies; Food chains and webs;
Geochemical cycles; Habitats and biomes; Lakes and limnology; Nutrient
cycles; Trophic levels and ecological niches.

Begon, M., J. L. Harper, and C. R. Townsend. Ecology: Individuals, Popula-
tions, and Communities. Oxford, England: Blackwell Scientific Publica-
tions, 1996.
Clark, Tim W., A. Peyton Curlee, Steven C. Minta, and Peter M. Kareiva,
eds. Carnivores in Ecosystems: The Yellowstone Experience. New Haven,
Colinvaux, Paul. Why Big Fierce Animals Are Rare: An Ecologist’s Perspective.
Princeton, N.J.: Princeton University Press, 1978.
189
Golly, Frank Benjamin. A History of the Ecosystem Concept in Ecology: More

than the Sum of the Parts. New Haven, Conn.: Yale University Press, 1993.
Hagen, Joel B. An Entangled Bank: The Origins of Ecosystem Ecology. New
Brunswick, N.J.: Rutgers University Press, 1992.
Hunter, Malcolm, Jr. Maintaining Biodiversity in Forest Ecosystems. New
York: Cambridge University Press, 1999.
Odum, Eugene P. Ecology and Our Endangered Life-Support Systems.
Real, Leslie A., and James H. Brown. Foundations of Ecology: Classic Papers
with Commentaries. Chicago: University of Chicago Press, 1991.
Smith, Robert Leo. Ecology and Field Biology. 6th ed. San Francisco:
Benjamin/Cummings, 2001.
190
ECOSYSTEMS: STUDIES
The study of ecosystems defines a specific area of the earth and the attendant inter-
actions among organisms and the physical-chemical environment present at the
site.
E cosystems are viewed by ecologists as basic units of the biosphere,

much as cells are considered by biologists to be the basic units of an or-
ganism. Ecosystems are self-organized and self-regulating entities within
which energy flows and resources are cycled in a coordinated, interdepen-
dent manner to sustain life. Disruptions and perturbations to, or within,
the unit’s organization or processes may reduce the quality of life there or
cause its demise. Ecosystem boundaries are usually defined by the re-
search or management questions being asked. An entire ocean can be
viewed as an ecosystem, as can a single tree, a rotting log, or a drop of pond
water. Systems with tangible boundaries—such as forests, grasslands,
ponds, lakes, watersheds, seas, or oceans—are especially useful to ecosys-
tem research.
Research Principles
The ecosystem concept was first put to use by American limnologist Ray-
mond L. Lindeman in the classic study he conducted on Cedar Bog Lake,
Minnesota, which resulted in his article “The Trophic Dynamic Aspect of
Ecology” (1942). Lindeman’s study, along with the publication of Eugene
P. Odum’s Fundamentals of Ecology (1953), converted the ecosystem notion
into a guiding paradigm for ecological studies, thus making it a concept of
theoretical and applied significance.
Ecologists study ecosystems as integrated components through which
energy flows and resources cycle. Although ecosystems can be divided into
many components, the four fundamental ones are abiotic (nonliving) re-
sources, producers, consumers, and decomposers. The ultimate sources of
energy come from outside the boundaries of the ecosystem (solar energy or
chemothermo energy from deep-ocean hydrothermal vent systems). Be-
cause this energy is captured and transformed into chemical energy by pro-
ducers and translocated through all biological systems via consumers and
decomposers, all organisms are considered as potential sources of energy.
Abiotic resources—water, carbon dioxide, nitrogen, oxygen, and other
inorganic nutrients and minerals—primarily come from within the bound-
191
Ecosystems: studies
aries of the ecosystem. From these, producers utilizing energy synthesize

biomolecules, which are transformed, upgraded, and degraded as they cy-
cle through the living systems that comprise the various components. The
destiny of these bioresources is to be degraded to their original abiotic
forms and be recycled.
The ecosystem approach to environmental research is a major endeavor.
It requires amassing large amounts of data relevant to the structure and
function of each component. These data are then integrated among the
components, in an attempt to determine linkages and relationships. This
holistic ecosystem approach to research involves the use of systems infor-
mation theory, predictive models, and computer application and simula-
tions. As ecosystem ecologist Frank B. Golley stated in his book A History
of the Ecosystem Concept in Ecology (1993), the ecosystem approach to the
study of ecosystems is “machine theory applied to nature.”
Research Projects
Initially, ecosystem ecologists used the principles of Arthur G. Tansley,
Lindeman, and Odum to determine and describe the flow of energy and
resources through organisms and their environment. Fundamental aca-
demic questions that plagued ecologists concerned controls on ecosystem
productivity: What are the connections between animal and plant produc-
tivity? How are energy and nutrients transformed and cycled in ecosys-
tems?
Once fundamental insights were obtained, computer-model-driven the-
ories were constructed to provide an understanding of the biochemo-
physical dynamics that govern ecosystems. Responses of ecosystem com-
ponents could then be examined by manipulating parameters within the
simulation model. Early development of the ecosystem concept culmi-
nated, during the 1960’s, in defining the approach of ecosystem studies.
Ecosystem projects were primarily funded under the umbrella of the In-
ternational Biological Program (IBP). Other funding came from the Atomic
Energy Commission and the National Science Foundation. The intention
of the IBP was to integrate data collected by teams of scientists at research
sites that were considered typical of wide regions. Although the IBP was
international in scope, studies in the United States received the greatest
portion of the funds—approximately $45 million during the life of IBP
(1964-1974).
Five major IBP ecosystem studies, involving grasslands, tundra, deserts,
coniferous forests, and deciduous forests, were undertaken. The Grass-
lands Project, directed by George Van Dyne, set the research stage for the
other four endeavors. However, because the research effort was so exten-
192
Ecosystems: studies
sive in scope, the objectives of the IBP were not totally realized. Because of
the large number of scientists involved, little coherence in results was ob-
tained even within the same project. A more pervasive concern, voiced by
environmentalists and scientists alike, was that little of the information ob-
tained from the ecosystem simulation models could be applied to the solu-
tion of existing environmental problems.
An unconventional project partially funded by the IBP was called the
Hubbard Brook Watershed Ecosystem. Located in New Hampshire and
studied by F. Herbert Bormann and Gene E. Likens, the project redirected
the research approach for studying ecosystems from the IBP computer-
model-driven theory to more conventional scientific methods of study.
Under the Hubbard Brook approach, an ecosystem phenomenon is ob-
served and noted. A pattern for the phenomenon’s behavior is then estab-
lished for observation, and questions are posed about the behavior. Hy-
potheses are developed to allow experimentation in an attempt to explain
the observed behavior. This approach requires detailed scrutiny of the eco-
system’s subsystems and their linkages. Since each ecosystem functions as
a unique entity, this approach has more utility. The end results provide in-
sights specific to the activities observed within particular ecosystems. Ex-
planations for these observed behaviors can then be made in terms of bio-
logical, chemical, or physical principles.
Utility of the Concept

Publicity from the massive ecosystem projects and the publication of Ra-
chel Carson’s classic Silent Spring (1962) helped stimulate the environmen-
tal movement of the 1960’s. The public began to realize that human activity
was destroying the bioecological matrices that sustained life. By the end of
the 1960’s, the applicability of the IBP approach to ecosystem research was
proving to be purely academic and provided few solutions to the problems
that plagued the environment. Scientists realized that, because of the lack
of fundamental knowledge about many of the systems and their links and
because of the technological shortcomings that existed, ecosystems could
not be divided into three to five components and analyzed by computer
simulation.
The more applied approach taken in the Hubbard Brook project, how-
ever, showed that the ecosystem approach to environmental studies could
be successful if the principles of the scientific method were used. The Hub-
bard Brook study area and the protocols used to study it were clearly de-
fined. This ecosystem allowed hypotheses to be generated and experimen-
tally tested. Applying the scientific method to the study of ecosystems had
practical utility for the management of natural resources and for testing
193
Ecosystems: studies
possible solutions to environmental problems. When perturbations such

as diseases, parasites, fire, deforestation, and urban and rural centers dis-
rupt ecosystems from within, this approach helps define potential mitiga-
tion and management plans. Similarly, external causative agents within
airsheds, drainage flows, or watersheds can be considered.
The principles and research approach of the ecosystem concept are be-
ing used to define and attack the impact of environmental changes caused
by humans. Such problems as human population growth, apportioning of
resources, toxification of biosphere, loss of biodiversity, global warming,
acid rain, atmospheric ozone depletion, land-use changes, and eutrophica-
tion are being holistically examined. Management programs related to
woodlands (the New Forestry program) and urban and rural centers (the
Urban to Rural Gradient Ecology, or URGE, program), as well as other gov-
ernmental agencies that are investigating water and land use, fisheries, en-
dangered species, and exotic species introductions, have found the ecosys-
tem perspective useful.
Ecosystems are also viewed as systems that provide the services neces-
sary to sustain life on earth. Most people either take these services for
granted or do not realize that such natural processes exist. Ecosystem re-
search has identified seventeen naturally occurring services, including
water purification, regulation, and supply, as well as atmospheric gas reg-
ulation and pollination. A 1997 article by Robert Costanza and others, “The
Value of the World’s Ecosystem Services and Natural Capital,” placed a
monetary cost to humanity should the service, for some disastrous reason,
need to be maintained by human technology. The amount is staggering,
averaging $33 trillion per year. Humanity could not afford this; the global
gross national product is only about $20 trillion.
Academically, ecosystem science has been shown to be a tool to dissect
environmental problems, but this has not been effectively demonstrated to
the public and private sectors, especially decision makers and policy-
makers at governmental levels. The idea that healthy ecosystems provide
socioeconomic benefits and services remains controversial. In order to
bridge this gap between academia and the public, Scott Collins of the Na-
tional Science Foundation suggested to the Association of Ecosystem Re-
search Centers that ecosystem scientists be “bilingual”; that is, they should
be able speak their scientific language and translate it so that the non-
scientist can understand.
Richard F. Modlin
See also: Biodiversity; Biomes: determinants; Biomes: types; Commu-

nities: ecosystem interactions; Desertification; Ecosystems: definition and
194
Ecosystems: studies
history; Geochemical cycles; Habitats and biomes; Hydrologic cycle; Nu-

trient cycles; Rain forests and the atmosphere; Trophic levels and ecologi-
cal niches.

Aber, J. D., and J. M. Melillo. Terrestrial Ecosystems. 2d ed. San Diego, Calif.:
Harcourt Academic Press, 2001.
Allen, T. F. H., and T. W. Hoekstra. Toward a Unified Ecology. New York: Co-
lumbia University Press, 1992.
Costanza, Robert, et al. “The Value of the World’s Ecosystem Services and
Natural Capital.” Nature 387, no. 6630 (May 15, 1997): 253-260.
Daily, Gretchen C., ed. Nature’s Services: Societal Dependence on Natural Eco-
systems. Washington, D.C.: Island Press, 1997.
Dodson, Stanley I., et al. Ecology. New York: Oxford University Press, 1998.
Golley, Frank B. A History of the Ecosystem Concept in Ecology. New Haven,
Likens, Gene E. The Ecosystem Approach: Its Use and Abuse. Oldendorf/
Luhe, Germany: Ecology Institute, 1992.
Vogt, Kristiina A., et al. Ecosystems: Balancing Science with Management.
New York: Springer, 1997.
195
ENDANGERED ANIMAL SPECIES
Endangered species are those varieties of plants and animals that are in immediate
danger of becoming extinct. Threatened species, by contrast, are those identified as
likely to become endangered in the near future.
A t the beginning of 2003, the U.S. Fish and Wildlife Service listed
nearly 400 animal species in the United States as “endangered”: in
immediate danger of extinction. Another 129 species of animals were des-
ignated as threatened, meaning likely to become endangered in the near
future. Globally, the World Conservation Union (IUCN) has placed thou-
sands of species on its list of endangered and vulnerable species, and more
are added every month. The International Council for Bird Preservation
has announced that more than one thousand of the nearly ten thousand
species of birds are endangered. Fish, especially freshwater fish, are among
the most immediately threatened types of animals. About one-fourth of
the worldwide number of species is in a state of dangerous decline.
Causes of Species Endangerment

The destruction of animal species is caused in four major ways: Humans
have hunted other species out of existence; habitats, the environments in
which plants or animals grow and develop, have been destroyed; new spe-
cies, such as rats, cats, goats, or ground-covering plants, have been intro-
duced into a region and displaced native species; and nonnative plants and
animals have introduced diseases into an environment, killing the existing
species. For much of history, hunting was the major cause of species extinc-
tion. However, hunting has become less of a factor because governments
and conservation authorities have imposed strict controls on the practice.
In the second half of the twentieth century, habitat destruction and inva-
sion by exotics (nonnative plants and animals) and the diseases they carry
caused the most damage. Most biologists agree that whatever the factors
involved, the rate of extinction has increased rapidly since the 1950’s.
Some people have argued that the destruction of a single species of fish,
bird, or flower would make little or no difference to the future of human
life or the earth. They also suggest that extinctions have always taken
place, even before human beings existed, and therefore are simply part of
the natural process of existence. Who, for instance, would really want to
have “saved” the dinosaurs? We should be glad they are gone. At least ten
196
Endangered animal species
million species exist, so who would miss a few dozen or a few hundred of
them?
These arguments ignore an important point. Each species inhabits a
small part of an entire ecosystem, a community of plants and animals that
are closely associated in a chain of survival. For example, plants absorb
chemicals and minerals from the soil that are essential to their health. Ani-
mals then eat the plants—grasses, fruits, leaves, or flowers—and digest the
nutrients they need for energy. Other animals, meat eaters (carnivores),
then eat these plant eaters and get their energy from them. If a single spe-
cies is removed from this chain, the whole ecosystem will experience con-
sequences that are difficult to predict and often negative or disastrous.
The death of an entire species constitutes a loss that cannot always be
measured in economic terms. The American biologist William Beebe made
the point that any species that is lost diminishes the quality of life for ev-
eryone:
The beauty and genius of a work of art may be reconceived, though its first
material expression can be destroyed; a vanished harmony may yet inspire
the composer, but when the last individual of a race of living things breathes
no more, another heaven and another earth must pass before such a one can
be again.
The bald eagle, and other bird populations, became endangered partly as a result of
the widespread use of DDT, a pesticide whose toxicity becomes concentrated as it
makes its way up the food chain. Agricultural runoff into streams and lakes causes
such pesticides to accumulate in the microorganisms on which fish feed, then the
fish, and finally the predators that eat the fish, from birds to humans. (PhotoDisc)
197
How Species Are Lost

The passenger pigeon is one example of species loss, a bird so numerous in
the 1820’s that John James Audubon, the famous American painter and
collector, wrote that the flapping of wings of flocks numbering in the hun-
dreds of millions on the Great Plains sounded like the roar of thunder.
More than nine billion of the pigeons were alive in 1850, yet slightly more
than sixty years later, exactly one bird, Martha, survived in the Cincinnati
Zoo, where she had been taken in 1912. The population fell from nine bil-
lion to one in little more than half a century, then to zero when Martha died
on September 1, 1917. People had found these pigeons delicious to eat and
easy to kill. They formed hundreds of hunting parties, killing more than
fifty thousand birds a week. No one dreamed the passenger pigeon could
ever be exterminated.
The same fate almost befell the American bison, often called the buffalo.
Before the coming of railroads and white settlers in the 1860’s and 1870’s,
the bison numbered more than 100 million. Native Americans hunted the
bison, eating their flesh and using the skins for clothing and shelter, but
Image Not Available
198
they killed only what they needed. The settlers, however, saw the bison as
a problem that needed to be solved. Huge herds of bison crossed railroad
tracks, forcing passenger trains to stop, and the animals interfered with
farming, knocking down fences and trampling grain fields. Railroad com-
panies and the U.S. Army sent out hunting parties to get rid of the bison.
By 1890 fewer than one thousand bison survived in a herd that had man-
aged to escape far into northern Canada. The extermination ended only af-
ter this small herd was given protection by the Canadian government.
Stories of other near extinctions are numerous and frightening but dem-
onstrate that action can be taken to save some if not all of the endangered
species. Whales, which had been hunted since the 1600’s, faced possible
extinction until action was taken to reduce hunting in the 1970’s. Whales
were easy to kill and provided oil and bone. Whale oil was the major sub-
stance burned in lamps until the electric light largely replaced oil-burning
lamps in the 1880’s. Europeans hunted the Atlantic whale, called the right
whale because it was the “right” one to kill, into virtual extinction by the
1860’s. When the right whale became too hard to find, hunters turned to
the Pacific right whale and then the bowhead whale before action was
taken by the world community to save remaining whales.
Greed and Ignorance

Human greed has brought death or near death to many species. The desire
for fur coats has nearly killed all jaguars, snow leopards, and various spe-
cies of fur seals. Pribilof Island fur seals were nearly hunted into extinction
in the late 1800’s. A treaty between the United States, Canada, and Russia
established limits on killing the species in its remote northern Pacific is-
land habitat, but enforcement has proved difficult, and thousands of seals
have been slaughtered despite international protection.
The belief that some animals are nuisances has led to the near extinction
of wolves, grizzly bears, cougars, and coyotes. These predators have been
poisoned and shot by the thousands and have become endangered species
as a result. Attempts to kill insects with pesticides in order to control the
spread of disease and improve crop yields were successful but had an un-
fortunate side effect. Chemicals from the pesticides worked their way into
ecosystems, killing millions of other forms of life. In the 1950’s, dichloro-
diphenyl-trichloroethane, or DDT, was used to kill malaria-carrying mos-
quitoes, but the chemical infested the whole food chain. It entered plants
that were eaten by animals and affected birds, fish, and butterflies. Pesti-
cide poisoning also diminished the numbers of the bald eagle and pere-
grine falcon, which started to come back only after rigid controls on pesti-
cides were established.
199
Events on the island of Madagascar, a large island in the Indian Ocean

off the east coast of Africa, demonstrate most fully the deadly conse-
quences of habitat destruction. About 180 million years ago, the island was
attached to the African continent, then was split off after a series of geologi-
cal catastrophes and ended up 250 miles to the east. The split occurred just
at the time mammals were emerging as a class of animals in Africa. One
mammal species, the monkeylike lemur, became isolated on Madagascar
and increased abundantly. Other animals caught on the island included
several kinds of giant birds, one weighing one thousand pounds and
standing ten feet tall. The island was isolated for millions of years, allow-
ing hundreds of species found nowhere else in the world to evolve in the
diverse island ecosystems. Madagascar had deserts, rain forests, dry for-
ests, and seashores. About 99 percent of its reptiles, 81 percent of its plants,
and 99 percent of its frogs were unique and tied specifically to the island’s
food chain. About nine thousand years ago, the Malagasy people began to
arrive on the island. They hunted, fished, began to grow crops, and in the
process destroyed more than 90 percent of the forests that covered Mada-
gascar. Dozens of species died as a result, including the giant elephant
bird, which was gone by 1700. Ten out of thirty-one species of lemur died
out by 1985. The loss of Madagascar’s forests caused terrible erosion,
which resulted in flooding and the destruction of more trees. Madagas-
car’s entire ecological system is now threatened, and hundreds of unique
species are listed as endangered. Only major restrictions on farming and
habitat destruction can save these animals.
Endangered Species Acts

The implications of species and habitat destruction were first described in
books such as Rachel Carson’s Silent Spring (1962). Carson, a biologist with
the U.S. Department of Interior, wrote about the effects of DDT and insecti-
cides on birds and other animals. Her book inspired Congress to pass the
Endangered Species Preservation Act in 1966. This law authorized the sec-
retary of the interior to protect certain fish and wildlife through the cre-
ation of a National Wildlife Refuge System. Congress strengthened the law
in 1969 by restricting importation of threatened species and adding more
domestic species to the list of those deserving protection.
The U.S. Department of the Interior published its first list of endangered
species in 1967. The list included seventy-two native species of animals, in-
cluding grizzly bears, certain butterflies, bats, crocodiles, and trout. No
plants were on this list. In 1973, President Richard M. Nixon signed into
law an Endangered Species Act that gave the secretaries of the interior and
of commerce responsibility for creating a list of endangered animals and
200
Endangered Animal Species

U.S. Foreign Total
Mammals 65 251 316
Birds 78 175 253
Reptiles 14 64 78
Amphibians 12 8 20
Fishes 71 11 82
Clams 62 2 64
Snails 21 1 22
Insects 35 4 39
Arachnids 12 0 12
Crustaceans 18 0 18
Total 388 516 904
Source: Data are from U.S. Fish & Wildlife Service, Threatened and Endangered Species
System (TESS), http://ecos.fws.gov/tess/html/boxscore.html, accessed on March 22,
2003.
plants, or species in immediate danger of extinction. Another list would

include species threatened with extinction, or those likely to become en-
dangered in the foreseeable future. Once an animal or plant was on ei-
ther of the lists, no one could kill, capture, or harm it. Penalties for viola-
tors were increased, and international or interstate trade of listed species
was prohibited. Fines up to ten thousand dollars could be imposed for
knowingly violating the act and one thousand dollars for unwittingly vio-
lating it.
A separate provision of the law mandated that federal agencies could
not engage in projects that would destroy or modify a habitat critical to the
survival of a threatened or endangered plant or animal. This provision be-
came a very important tool in the battle to save species. Friends of wildlife
used it to block highway and dam projects, at least until government offi-
cials could prove that construction would have no major impact on a frag-
ile ecosystem. The law even called for affirmative measures to aid in the re-
covery of listed species. The secretaries of the interior and of commerce
were required to produce recovery plans detailing steps necessary to bring
a species back to a point where it no longer needed protection. Money was
appropriated for states to design recovery programs, and most states es-
tablished plans of their own to deal with local crises.
201
A major threat to the 1973 law arose in 1978 during the Tellico Dam con-
troversy. The Tennessee Valley Authority, a federal government agency,
proposed building a hydroelectric dam on the Tennessee River in Loudon
County, Tennessee, in the late 1970’s. Shortly after plans were made public,
a scientist from the University of Tennessee discovered a three-inch-long
fish, the snail darter, that was unique to that area. Building the dam, a $250
million project, would destroy that snail darter’s habitat and eliminate the
fish. Environmentalists successfully argued in federal court that the dam
had to be abandoned. The U.S. Supreme Court supported the ruling of the
lower court, arguing that when Congress passed the law, it had intended
that endangered species be given the highest priority regardless of the cost
or other concerns involved. However, in 1981, Congress enacted a special
exemption that excluded “economically important” federal projects from
the act. A federal judge then found the Tellico Dam to be without economic
importance, and construction was again halted. Congressmen friendly to
dam interests then slipped an amendment directing completion of the dam
onto an unrelated environmental bill, which passed, and Tellico was con-
structed. However, the principle of protection remained intact, and the
power of the 1973 act remained in force. The snail darter apparently sur-
vived, too, as scientists found it living in a river not far from the spot where
it was originally discovered.
International Trade Bans

The 1973 act also made the United States a partner in the Convention on In-
ternational Trade in Endangered Species of Wild Flora and Fauna. This
treaty came out of a conference in Washington, D.C., attended by represen-
tatives from eighty nations. It created an international system for control of
trade in endangered species. Enforced by the World Conservation Union
(IUCN) headquartered in Switzerland, the convention has more than one
hundred members. The IUCN publishes a series of lists in its Red Data Book
designating three categories of species. Category one consists of those in
immediate danger and therefore absolutely banned from international
hunting and trading. Animals and plants in categories two and three are
not immediately threatened but require special export permits before they
can be bought and sold because their numbers have been seriously re-
duced.
This convention has a major flaw, a loophole that can be exploited by
any member. A nation can make a “reservation” on any listed species, ex-
empting itself from the ban on trade. Japan has been the most frequent user
of the reservation, exempting itself from controls on the fin whale, the
hawksbill turtle, and the saltwater crocodile, all on the IUCN’s most en-
202
dangered list. Unless this loophole is closed, the IUCN can do little to save
extremely threatened species.
Restoring Endangered Species

Several species in the United States—the California condor, the black-
footed ferret, whooping cranes, and a bird called the Guam rail—have
been saved from extinction because of the 1973 Endangered Species Act.
The road to extinction has also been reversed for the brown pelican, found
in the southeastern states, the American alligator, which had almost been
hunted to extinction in Florida, and the perigrine falcon in the eastern
states. Other species, however, such as the dusky seaside sparrow and the
Palos Verdes butterfly, have totally disappeared.
The spotted owl found in parts of the rain forest in Oregon and Wash-
ington has attracted a good deal of attention because of efforts to save it.
The case of the owl points to the most difficult questions raised by the act:
Which comes first, the welfare of the plant or animal, or the economic
needs of people? Each pair of spotted owls needs six to ten square miles of
forest more than 250 years old in which to hunt and breed. The owls also
need large hollow trees for nesting, as well as large open fields in which to
search for mice and other small animals. A suitable ecosystem that meets
all these needs is found only in parts of twelve national forests in the re-
gion. At the same time, loggers in the areas need jobs. When the two inter-
ests, represented by the lumber industry and environmentalists, collide, it
is left to the courts to determine which interest will prevail or whether a
compromise can be arranged.
Outside the United States, the future of endangered species appears
much grimmer. Scientists at IUCN think several hundred thousand species
will disappear by the end of the second decade of the twenty-first century.
Many of these species have never even been identified or named. The most
endangered habitats in the world are the tropical rain forests, which were
reduced by half, nearly 3.5 million square miles, during the twentieth cen-
tury. About 43,000 square miles are destroyed each year, mainly to provide
farms and cattle ranches. The most threatened large animal species in these
forests are the large cats, including tigers, jaguars, and leopards; fifteen of
the twenty-five species of cats that live in the forests are on the most endan-
gered list.
One solution to forest destruction has been the creation of large wildlife
refuges. Several African nations have created one or more of these, but
there are limits to the amount of land available for conservation efforts.
Another solution is the establishment of more wildlife zoos. Several zoos
have successful programs for saving species on the very edge of extinction.
203
However, capacity is limited, and the very small numbers of animals in a

zoo’s herd create problems of interbreeding and the handing down of re-
cessive genes.
For many species, it is too late to do very much, so scientists and biolo-
gists divide populations into three groups: those that can survive without
help, those that would die regardless of whether help is provided, and
those species that might survive with help and would certainly die with-
out it. Environmentalisms and restoration ecologists focus their efforts on
the plants and animals that fall into the third category. Resources are lim-
ited, however, and much work needs to be done, or extinctions will take
place at a pace as yet unseen in the history of living things.
Leslie V. Tischauser
See also: Balance of nature; Biodiversity; Communities: ecosystem interac-

tions; Conservation biology; Deforestation; Ecosystems: definition and
history; Ecosystems: studies; Endangered plant species; Extinctions and
evolutionary explosions; Genetic diversity; Habitats and biomes; Nonran-
dom mating, genetic drift, and mutation; Old-growth forests; Pollution ef-
fects; Reefs; Reforestation; Restoration ecology; Species loss; Sustainable
development; Trophic levels and ecological niches; Urban and suburban
wildlife; Wildlife management; Zoos.

Beacham, Walton, and Kirk H. Beets, eds. Beacham’s Guide to International
Endangered Species. Osprey, Fla.: Beacham, 1998.
Disilvestro, Roger L. The Endangered Kingdom: The Struggle to Save America’s
Wildlife. New York: John Wiley & Sons, 1989.
Endangered Species of the World. 2 vols. Detroit: Gale Research, 1994.
Endangered Wildlife of the World. 11 vols. New York: Marshall Cavendish,
1993.
Sherry, Clifford J. Endangered Species: A Reference Handbook. Santa Barbara,
Calif.: ABC-Clio, 1998.
Wilson, Edward O. The Diversity of Life. New York: W. W. Norton, 1993.
204
ENDANGERED PLANT SPECIES
The World Conservation Union defines “endangered species” as those in immedi-

ate danger of extinction and “threatened species” as those at a high risk of extinc-
tion but not yet endangered. Vulnerable species are considered ones that are likely
to become extinct at some point in the foreseeable future. Rare species are at risk
but not yet at the vulnerable, threatened, or endangered levels.
W orldwide, the number of endangered plant species was estimated at

more than 33,418 in 1999. This number is much higher than that of
all of the endangered or threatened animal species combined. Although
extinction is a natural process, and all species will eventually be extinct,
human activities threaten the existence of plant and animal life worldwide.
Humans use plants for food; medicine; building materials; energy; to
clean water, air, and soil of pollutants; to control erosion; and to convert
carbon dioxide to oxygen. The process of extinction increased dramatically
during the nineteenth and twentieth centuries because of habitat destruc-
tion or loss, deforestation, competition from introduced species, pollution,
global warming, and plant hunting, collecting, and harvesting. Over time,
pollutants and contaminants accumulate in the soil and remain in the envi-
ronment, some for many decades. Pollution in the atmosphere also con-
tributes to long-term changes in climate.
Habitat loss
By far the most significant threat to plant species is habitat loss or destruc-
tion. Habitat loss can occur because of resource harvesting for food, medi-
cine, and other products; deforestation; and the conversion of wilderness
for agricultural, industrial, or urban uses. Wood consumption and tree
clearing for agriculture and development threaten the world’s forests, es-
pecially the tropical forests, which may disappear by the mid-twenty-first
century if sufficient preventive action is not taken. Natural disasters, such
as climatic changes, meteorites, floods, volcanic eruptions, earthquakes,
hurricanes, drought, and tornados, also can be devastating to a habitat.
In Europe and Asia, the plant distribution is complex, with isolated
populations of plants spread across a large area. The plants are greatly
influenced by the cold climate and by humans. Plant species are disap-
pearing, especially in Europe and the Mediterranean, because of habitat
destruction and disturbances including urbanization, road construction,
205
Endangered plant species
Image Not Available
overgrazing, cultivation, forest plantation, fire, pollution, and overexploi-

tation of resources or for use in horticulture.
Mountain plants are threatened by development for industry and tour-
ism, pollution, strip mining, walkers, and skiers. The wetlands are threat-
ened by removal of peat for fuel, water extraction which lowers the water
table, and increased drainage for building or agriculture or fear of malaria.
Recreational use and susceptibility to pollution such as acid rain or fertil-
izer run-off present further threats.
In North America, the major causes of endangerment include loss of
habitat, overexploitation of resources, introduction of invasive species,
and pollution. A massive loss of wilderness has occurred through the
clearing of forests, plowing of prairies, and draining of swamplands. For
example, in the northeastern United States there are only 13 square miles of
alpine habitat, an area in which grow thirty-three at-risk species. This area
is heavily used by hikers and mountain bikers. The Florida Everglades are
threatened because the water supply is diverted to supply cities, industry,
and agriculture.
In California, the Channel Islands are home to seventy-six flowering
plants which do not exist on the mainland. Eighteen species are located on
206
just one island. These plants, including the San Clemente broom, bush
mallow (Malacothamnus Greene), a species of larkspur, and the San Cle-
mente Island Indian paintbrush (Castilleja grisea), have been devastated by
introduced grazers, browsers, and by invasive other plants. In Hawaii,
more than 90 percent of native plants and almost all land birds and inverte-
brates are found nowhere else in world. The Hawaiian red-flowered gera-
nium Geranium arboreum is threatened by introduced feral pigs, agricul-
tural livestock, and competition by nonnative plants.
In developing or highly populated nations in Asia, Africa, Central and
South America, the Caribbean, the Pacific Ocean islands, Australia, and
New Zealand, habitat loss occurs because of population needs. Land is
cleared for agriculture, development, and population resettlement. In Cen-
tral America and the Caribbean, the Swietenia mahogany is found only in
a few protected or remote areas. The Caoba tree (Persea theobromifolia) was
newly identified as a species as recently as 1977. The lumber is commer-
cially important, and habitat loss has occurred due to the conversion of for-
ests to banana and palm plantations. In Ecuador, only 6 percent of the orig-
inal rain forest remains standing, because the rest has been converted to
farmland. In Asia, including the Philippines, population pressures bring
about deforestation and the clearing of land for agriculture.
In southern Africa, land is used for crops, livestock, and firewood pro-
duction. Overgrazing and the introduction of agriculture have caused the
Sahara Desert area to grow rapidly. The island of Madagascar has between
ten thousand and twelve thousand plant species, of which 80 percent grow
nowhere else in the world. Because of conversion to grassland through
farming methods, only about one-fifth of the original species survive. In
Australia there are 1,140 rare or threatened plants, and logging, clearing
for grazing animals and crops, building developments, and mining have
threatened many native species.
Plant Hunting, Collecting, and Harvesting

Habitat damage, the construction of facilities, and the opening of remote
areas for human population have made many plants vulnerable to gather-
ing and collecting. Some plants have been overharvested by gardeners,
botanists, and horticulturists. One species of lady’s slipper orchid (Cypri-
pedium calceolus) is rare over much of its natural range except in parts of
Scandinavia and the Alps because of collecting. Additionally, many moun-
tain flowers or bulbs such as saxifrages, bellflowers, snowdrops, and cycla-
men are endangered. In France and Italy, florulent saxifrage (Saxifraga
florulenta), an alpine plant, has been overcollected by horticulturists and
poachers.
207
Parts of the southeastern United States have poor soil that is home to the
carnivorous or insectivorous plants—those that eat insects. These plants
include sundews, bladderworts, Venus’s flytrap, and pitcher plants. Col-
lectors or suppliers have stripped many areas of all of these plants. In the
Southwest, rare cacti are harvested for sale nationwide and worldwide.
Endangered cacti include the Nellie Cory cactus (which has one remaining
colony), Epithelantha micromeres bokei, Ancistrocactus tubuschii, saguaro cac-
tus (Carnegiea gigantea), and Coryphantha minima. Near the Sierra Madre,
two tree species—Guatemalan fir, or Pinabete, and the Ayuque—are en-
dangered because of harvesting for use as Christmas trees or for the mak-
ing of hand looms. Additionally, sheep eat the seedlings. In New Mexico,
the gypsum wild buckwheat habitat is limited to one limestone hill, and
the plants are threatened by cattle, off-road vehicles, and botanists.
In southern Mexico, there are 411 species of epiphytes (air plants or bro-
meliads in the genus Tillandsia), of which several are extremely rare. These
plants are threatened by overcollection for the houseplant trade or conser-
vatories. The African violet (Saintpaulia ionantha) of Tanzania may soon be
extinct in the wild because of the horticultural trade and habitat loss due to
encroaching agriculture.
Worldwide, orchids are overcollected for horticulture. Several species
have been collected to extinction, are extremely rare, or have been lost be-
cause of habitat destruction. Examples include the extremely rare blue
vanda (Vanda caerulea); Paphiopedilum druryi, believed extinct in its native
habitat; Dendrobium pauciflorum, endangered and possibly extinct—only a
single plant was known to exist in the wild in 1970; and the Javan phalae-
nopsis orchid, Phalaenopsis javanica. The latter was believed extinct. When
Endangered Plant Species

U.S. Foreign Total
Flowering plants 570 1 571
Conifers and cycads 2 0 2
Ferns and allies 24 0 24
Lichens 2 0 2
Totals 598 1 599
Source: Data are from U.S. Fish & Wildlife Service, Threatened and Endangered Species
System (TESS), http://ecos.fws.gov/tess/html/boxscore.html, accessed on March 22,
2003.
208
it was rediscovered in 1960’s, it was overcollected by commercial orchid

dealers and thereby exterminated. There are no other known wild popula-
tions.
About 80 percent of the human populations in developing countries
rely on traditional medicine, of which 85 percent of ingredients come from
plant extracts. In Western medicine, one in four prescription medicines
contain one or more plant products. Some at-risk species contain chemicals
used in treating medical conditions, such as the African Prunus africana
tree, whose bark has chemicals used to treat some prostate gland condi-
tions, and the Strophanthus thollonii, a root parasite with chemicals used in
heart drugs.
The Madagascan periwinkle (Catharanthus roseus) is commonly culti-
vated (its close relative Catharanthus coriaceus is rare and its medicinal im-
portance unknown) and produces about seventy chemicals, some of which
are useful in the treatment of cancer. The Indian podophyllum (Podo-
phyllum hexandrum), a threatened species, is used to treat intestinal worms,
constipation, and cancer. Rauwolfia (Rauvolfia serpentina), also a threat-
ened species, is used to treat mental disorders, hypertension, and as a seda-
tive. The lily Amorphopahllus campanulatus is used to treat stomachaches,
and a fig, Ficus sceptica, is used to treat fever; both of these species are vul-
nerable because of habitat destruction.
The Micronesian dragon tree is believed to have magical and medicinal
properties. It has been overharvested and is now extinct on several islands.
In the United States’ Appalachian Mountains, American ginseng is being
overcollected because of an escalating demand for this plant’s health bene-
fits.
Conservation
The conservation of endangered plant species employs several compelling
arguments: Plants enhance the world’s beauty, have the right to exist, and
are useful to people. The most persuasive argument may be that the sur-
vival of the human species depends on a healthy worldwide ecosystem.
Three major goals of conservation are recovery, protection, and reintroduc-
tion.
Conservation methods depend on increasing public awareness by pro-
viding information about endangered or threatened species so that people
can take action to reverse damage to the ecosystems. Other important
strategies include achieving a widespread commitment to conservation
and obtaining funding to protect rare or endangered species. Conservation
efforts include setting aside protected areas, such as reserves, wilderness
areas, and parks, and recognizing that humans must integrate and protect
209
biodiversity where they live and work. Many countries are actively con-
serving species through protected areas, endangered-species acts, detailed
studies of species and habitat, and information campaigns directed to the
public.
Virginia L. Hodges
See also: Balance of nature; Biodiversity; Communities: ecosystem interac-

tions; Conservation biology; Deforestation; Ecosystems: definition and
history; Ecosystems: studies; Endangered animal species; Extinctions and
evolutionary explosions; Genetic diversity; Habitats and biomes; Nonran-
dom mating, genetic drift, and mutation; Old-growth forests; Pollution ef-
fects; Reefs; Reforestation; Restoration ecology; Species loss; Sustainable
development; Trophic levels and ecological niches; Urban and suburban
wildlife; Wildlife management; Zoos.

Burton, John A., ed. Atlas of Endangered Species. New York: Macmillan,
1999.
Crawford, Mark. Habitats and Ecosystems: An Encyclopedia of Endangered
America. Santa Barbara, Calif.: ABC-CLIO, 1999.
Freedman, Bill, ed. Encyclopedia of Endangered Species. Vol. 2. Detroit: Gale
Research, 1999.
Wilson, Edward O. The Future of Life. New York: Alfred A. Knopf, 2001.
210
EROSION AND EROSION
CONTROL
Types of ecology: Agricultural ecology; Ecosystem ecology; Restoration

and conservation ecology; Soil ecology
Erosion is the loss of topsoil through the action of wind and water. Erosion control
is vital because soil loss from agricultural land is a major contributor to nonpoint-
source pollution and desertification and represents one of the most serious threats
to world food security.
I n the United States alone some two billion tons of soil erode from crop-
land on an annual basis. About 60 percent, or 1.2 billion tons, is lost
through water erosion, while the remainder is lost through wind erosion.
This is equivalent to losing 0.3 meter (1 foot) of topsoil from two million
acres of cropland each year. Although soil is a renewable resource, soil for-
mation occurs at rates of just a few inches per hundred years, which is
much too slow to keep up with erosive forces. The loss of soil fertility is in-
calculable, as are the secondary effects of polluting surrounding waters
and increasing sedimentation in rivers and streams.
Erosion removes the topsoil, the most productive soil zone for crop pro-
duction and the plant nutrients it contains. Erosion thins the soil profile,
which decreases a plant’s rooting zone in shallow soils, and can disturb the
topography of cropland sufficiently to impede farm equipment operation.
It carries nitrates, phosphates, herbicides, pesticides, and other agricul-
tural chemicals into surrounding waters, where they contribute to cultural
eutrophication. Erosion causes sedimentation in lakes, reservoirs, and
streams, which eventually require dredging.
Water Erosion
The common steps in water erosion are detachment, transport, and deposi-
tion. Detachment releases soil particles from soil aggregates, transport car-
ries the soil particles away and, in the process, scours new soil particles
from aggregates. Finally, the soil particles are deposited when water flow
slows. In splash erosion, raindrops impacting the soil can detach soil parti-
cles and hurl them considerable distances. In sheet erosion, a thin layer of
soil is removed by tiny streams of water moving down gentle slopes. This
is one of the most insidious forms of erosion because the effects of soil loss
are imperceptible in the short term. Rill erosion is much more obvious be-
211
Erosion and erosion control
cause small channels form on a slope. These small channels can be filled in
by tillage. In contrast, ephemeral gullies are larger rills that cannot be filled
by tillage. Gully erosion is the most dramatic type of water erosion. It
leaves channels so deep that even equipment operation is prevented. Gully
erosion typically begins at the bottom of slopes where the water flow is
fastest and works its way with time to the top of a slope as more erosion oc-
curs.
Wind Erosion
Wind erosion generally accounts for less soil loss than water erosion, but in
states such as Arizona, Colorado, Nevada, New Mexico, and Wyoming, it
is actually the dominant type of erosion. Wind speeds 0.3 meter (1 foot)
above the soil that exceed 16 to 21 kilometers per hour (10 to 13 miles per
hour) can detach soil particles. These particles, typically fine- to medium-
size sand fewer than 0.5 millimeter (0.02 inch) in diameter, begin rolling
and then bouncing along the soil, progressively detaching more and more
soil particles by impact. The process, called saltation, is responsible for 50
to 70 percent of all wind erosion. Larger soil particles are too big to become
suspended and continue to roll along the soil. Their movement is called
surface creep.
Topsoil erosion is one of the most economically devastating forms of erosion, caused
not only by wind and water erosion but also by human disturbance resulting from
agriculture, overgrazing, deforestation, and soil compaction. (PhotoDisc)
212
The most obvious display of wind erosion is called suspension, when

very fine silt and clay particles detached by saltation are knocked into the
air and carried for enormous distances. The Dust Bowl of the 1930’s was
caused by suspended silt and clay in the Great Plains of the United States.
It is also possible to see the effects of wind erosion on the downward side of
fences and similar obstacles. Wind passing over these obstacles deposits
the soil particles it carries. Other effects of wind erosion are tattering of
leaves, filling of road and drainage ditches, wearing of paint, and increas-
ing incidence of respiratory ailments.
Erosion Control
The four most important factors affecting erosion are soil texture and struc-
ture, roughness of the soil surface, slope steepness and length, and soil
cover. There are several passive and active methods of erosion control that
involve these four factors. Wind erosion, for example, is controlled by cre-
ating windbreaks, rows of trees or shrubs that shorten a field and reduce
the wind velocity by about 50 percent. Tillage perpendicular to the wind
direction is also a beneficial practice, as is keeping the soil covered by plant
residue as much as possible.
Water erosion is controlled by similar cultural practices. For example,
highly erosive, steeply sloped land can be protected by placing it in the
U.S.-government-sponsored Conservation Reserve Program. Tillage can
be done along the contour of slopes. Long slopes can be shortened by ter-
racing, which also reduces the slope steepness. Permanent grass water-
ways can be planted in areas of cropland that are prone to water flow. Like-
wise, grass filter strips can be planted between cropland and adjacent
waterways to impede the velocity of surface runoff and cause suspended
soil particles to sediment and infiltrate before they can become contami-
nants.
Conservation tillage practices, such as minimal tillage and no-tillage,
are being widely adapted by farmers as a simple means of erosion control.
As the names imply, these are tillage practices in which as little disruption
of the soil as possible occurs and in which any crop residue remaining after
harvest is left on the soil surface to protect the soil from the impact of rain
and wind. The surface residue also effectively impedes water flow, which
causes less suspension of soil particles. Because the soil is not disturbed,
practices such as no-tillage also promote rapid water infiltration, which
also reduces surface runoff. No-tillage is rapidly becoming the predomi-
nant tillage practice in southeastern states such as Kentucky and Tennes-
see, where high rainfall and erodible soils occur.
Mark S. Coyne
213
See also: Desertification; Grazing and overgrazing; Integrated pest man-

agement; Multiple-use approach; Rangeland; Reforestation; Slash-and-
burn agriculture; Soil; Soil contamination.

Gershuny, Grace, and Joseph Smillie. The Soul of Soil: A Guide to Ecological
Soil Management. 3d ed. Davis, Calif.: agAccess, 1995.
Morgan, R. P. C. Soil Erosion and Conservation. New York: Wiley, 1995.
Plaster, Edward. Soil Science and Management. Albany, N.Y.: Delmar, 1997.
Schwab, Glenn O., et al., eds. Soil and Water Conservation Engineering. 4th
ed. New York: Wiley, 1993.
214
ETHOLOGY
Ethology is the study of animal behavior from the perspective of zoology. The infor-
mation acquired through ethology has helped scientists better understand animals
in all their variety.
E thology is the branch of zoology that investigates the behavior of ani-

mals. Behavior may be defined as all the observable responses an ani-
mal makes to internal or external stimuli. Responses may be either move-
ments or secretions; however, the study of behavior is much more than a
descriptive account of what an animal does in response to particular stimuli.
The ethologist is interested in the “how” and “why” of the behaviors they
observe. Answering such questions requires an understanding of the phys-
iology and ecology of the species studied. Those who study animal behavior
are also interested in the ultimate or evolutionary factors affecting behavior.
The Roots of Ethology

Ethology is a young science, yet it is also a science with a long history. Prior
to the late nineteenth century, naturalists had accumulated an abundance
of information about the behavior of animals. This knowledge, although
interesting, lacked a theoretical framework. In 1859, Charles Darwin pub-
lished On the Origin of Species by Means of Natural Selection, and with it pro-
vided a perspective for the scientific study of behavior. Behavior was more
central to two of Darwin’s later books, The Descent of Man and Selection in
Relation to Sex (1871) and Expression of the Emotions of Man and Animals
(1873). By 1973, the science of ethology was sufficiently well developed to
be acknowledged by the presentation of the Nobel Prize for Physiology or
Medicine to Nikolaas Tinbergen, Konrad Lorenz, and Karl von Frisch for
their contributions to the study of behavior. The work of these men was
central to the development of modern ethology.
The experimental studies of Frisch revealed the dance language of the
honeybee and ways in which the sensory perception of the bees differs
from the human sensory world. An awareness of species-specific sensory
abilities has provided an important research area and has emphasized a
factor that must be considered in the experimental design and interpreta-
tion of many types of behavioral research.
Tinbergen studied behavior in a variety of vertebrate and invertebrate
organisms. He was good both at observation of animals in their natural
215
Ethology
habitat and in the design of simple but elegant experiments. His 1951 book
The Study of Instinct is a classic synthesis of the knowledge that had been
gained through the scientific study of animal behavior of that time.
Konrad Lorenz is considered by many to be the founder of ethology, be-
cause he discovered and effectively publicized many of the classic phe-
nomena of ethology. Pictures of Lorenz being followed by goslings are al-
most a standard feature of texts that discuss the specialized form of
learning known as imprinting. In natural settings, imprinting allows young
animals to identify their parents appropriately. Another contribution of
Lorenz was his book King Solomon’s Ring: New Light on Animal Ways, pub-
lished in 1952. This extremely readable book raised public awareness of the
scientific study of animal behavior and kindled the interest of many who
eventually joined the ranks of ethologists.
Ethology and Neurobiology

Many of the features of ethological research characteristic of the work of
Lorenz, Tinbergen, and von Frisch have continued to be characteristic of
the field. They were concerned that the behavior of animals be understood
in the context of the species’ natural habitat and that both proximate and
ultimate levels of explanation would be examined. Their research strate-
gies have been supplemented by an increase in laboratory-based research
and by the introduction of new types of experimental design. These devel-
opments have softened the distinctions between ethology and another
field of behavioral study, comparative psychology. The focus of compara-
tive psychology is comparative studies of the behavior of nonhuman ani-
mals. Initially, questions about learning and development were the major
problems investigated in comparative psychology. Although the animals
most frequently studied were primates and rodents, those doing the re-
search were interested in gaining insight into the behavior of humans.
Comparative psychology was long dominated by behaviorism, a school
of thought that assumes that the ultimate basis of behavior is learning.
The behaviorists employed rigorous experimental methods. Because such
methods require carefully controlled conditions, behavioral research is
typically laboratory based, and animals are therefore tested in surround-
ings remote from their natural environment. Over time, comparative psy-
chology has broadened both the questions it asks and the organisms it
studies. The boundaries between comparative psychology and ethology
have been further blurred by the rising number of scientists crossing disci-
plinary lines in their research. Each discipline has learned from the other,
and both have also profited from knowledge introduced through neuro-
biology and behavioral genetics.
216
Ethology
Neurobiology investigates the structure and function of the nervous

system. One area of ethology that has been directly enriched through
neurobiology is the study of sensory perception in animals. The techniques
developed in neurobiology allow the investigator to record the response of
many individual neurons simultaneously. The neurobiologist examines
phenomena such as stimulus filtering at the level of the cell. Stimulus filter-
ing refers to the ability of nerve cells to be selective in their response to stim-
uli. For example, moths are highly sensitive to sounds in the pitch range of
sounds made by the bats that are their chief predators. Neurobiology pro-
vides a powerful tool for understanding behavior at the proximate level.
Behavioral Genetics
Another source of information for the ethologist is behavioral genetics.
Because of the evolutionary context of ethology, it is important to have an
understanding of the genetic basis of behavior. If there were no genetic
component in behavior, behavior would not be subject to natural selection.
(Natural selection refers to the process by which some genes increase in fre-
quency in a population while alternates decrease because the favored genes
have contributed to the reproductive success of those organisms that have
them.) While the ethological approach to behavior assumes that behavior
patterns are the result of interactions between genes and environment, in-
vestigators often ask questions about the genetic programming of behavior.
Early ethologists performed isolation and cross-fostering experiments
to discover whether behaviors are learned or instinctive. If a behavior ap-
pears in an individual that has been reared in isolation without the oppor-
tunity to learn, the behavior is considered instinctive. Observing the be-
havior of an individual reared by parents of a different species is similarly
revealing. When behavior patterns of conspecifics appear in such cross-
fostered individuals, such behaviors are regarded as instinctive. Instinc-
tive behaviors typically are innate behaviors that are important for sur-
vival. For example, one very common instinctive behavior is the begging
call of a newly hatched bird. Isolation and cross-fostering experiments are
still a part of the experimental repertoire of ethologists, but behavioral ge-
netics permits the asking of more complex questions. For example, a be-
havior may accurately be labeled instinctive, but it is more revealing to de-
termine the developmental and physiological processes linking a gene or
genes to the instinctive behavior.
Behavioral Ecology
The ethologist is also interested in determining whether behavior is adap-
tive. It is not sufficient to identify what seems to be a commonsense advan-
217
Ethology
tage of the behavior. It is important to show that the behavior does in

fact contribute to reproductive success in those that practice the behavior
and that the reasons the behavior is adaptive are those that are hypothe-
sized. When behaviors are tested, they frequently do turn out to be adap-
tive in the ways hypothesized. This type of research, however, has pro-
vided many surprises. Research on the adaptive value of behaviors in
coping with environmental problems that affect reproductive success is
known as behavioral ecology, a major subdiscipline of both ethology and
ecology.
Behavioral ecology addresses a variety of questions, in part because the
process of evolution is opportunistic. For any environmental problem
there are alternate solutions, and the solution a particular species adopts is
dependent upon the possibilities inherent in its genes. Questions ad-
dressed include such things as whether a species is using the optimum
strategy or how a species benefits from living in a group. Because alternate
strategies are possible even within a species, behavioral ecologists are in-
terested in evolutionarily stable strategies. An evolutionarily stable strat-
egy is a set of behavioral rules that, when used by a particular proportion
of a population, cannot be replaced by any alternative strategy. For exam-
ple, the sex ratio present in a particular population will determine the opti-
mum sex ratio for the offspring of any individual.
Sociobiology
Sociobiology is another major area of modern ethology. Sociobiology ex-
amines animal social behavior within the framework of evolution. Animal
species vary in the degree of social behavior they exhibit; other variables
include group size and the amount of coordination of activities occurring
within the group. The sociobiologist is interested in a number of questions,
but prominent among them are the reasons for grouping. Hypotheses such
as defense against predators or facilitation of reproduction can be tested.
The particular advantage or advantages gained by grouping varies among
species. Two important concepts in sociobiology are kin selection and in-
clusive fitness.
Kin selection refers to the differential reproduction of genes that affect
the survival of offspring or closely related kin. Behavior such as the broken-
wing display of the killdeer is an example. The behavior carries risk but
would be promoted by selection if the offspring of individuals using the
display were protected from predators often enough to compensate for the
risk. Inclusive fitness is the term used to recognize the concept that fitness
includes the total genotype, including those genes that may lower the indi-
vidual’s survival as the price of leaving more genes in surviving kin. The
218
Ethology
concepts of kin selection and inclusive fitness help to address one problem
raised by Darwin, the question of altruistic behavior. Ethology is a young
science but a very exciting one, because there are so many questions that
can be asked about animal behavior within the context of evolution.
Studying Ethology
The methods and tools of ethology cover the entire spectrum of complex-
ity. One simple, but demanding, method is to collect normative data about
a species. In its simplest form, the scientist observes what an animal does
and writes it down in a field notebook. Finding and following the animal,
coping with field conditions such as bad weather and rugged terrain, and
keeping field equipment in operating condition add challenge and variety
to this approach. The ethologist uses various techniques to get data as un-
biased as possible. One of these is to choose a focal animal at random (or on
a rotation) and observe the focal animal for a specific amount of time be-
fore switching observation to another member of the population. This pre-
vents bias in which individuals and which behaviors are observed. The
sampling of an individual’s behavior at timed intervals is an even more ef-
fective way of avoiding bias.
When all or most of an animal’s behavioral repertoire is known, a list
known as an ethogram can be constructed. This catalog can be organized
into appropriate categories based on function. Ethograms provide useful
baseline information about the behavior of a species. For animals that are
difficult or impossible to follow, radio-tracking techniques have been de-
veloped. Collars that emit radio signals have been designed for many ani-
mals. Miniaturization has made it possible for radio tracking to be used
even on relatively small animals.
In field studies, animals are often marked in some way so that observers
are able to follow individual animals. A number of techniques have been
developed, including banding birds with colored acrylic bands. Color
combinations can be varied so that each member of the population has
a unique combination. Marking allows the observer to get information
such as individual territory boundaries and to determine which animals
interact.
Models are frequently used in experiments. For example, a model can
be used to determine whether individuals in a species need to learn to
identify certain classes of predators. Models were used in many of the clas-
sic experiments in ethology. Modern technology has allowed the develop-
ment of much more sophisticated models. One of the most interesting is a
“bee” that can perform a waggle dance (used by bees to indicate location)
so effectively that its hivemates can find the food source. Whether a model
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Ethology
is simple or sophisticated, it can provide a tool to determine the cues that

trigger an animal’s response.
Neurobiologists use electrodes and appropriate equipment to stimulate
and record the responses of neurons. They can also stimulate specific re-
gions of the nervous system by using tiny tubes to deliver hormones or
neurotransmitters. Genetic technology has made it possible to examine the
deoxyribonucleic acid (DNA) of individuals in a species. This tool can be
used, for example, to determine whether females in monogamous species
are completely monogamous or whether some of their offspring are fa-
thered by males other than their mates.
Tape recorders have become very important in studies of animal vocal-
izations. Recorders are used in two ways. The animal’s vocalizations may
be recorded and the recording used to make sound spectrographs for anal-
ysis. The recordings may also be used to determine whether individuals
can discriminate between the vocalizations of neighboring and nonneigh-
boring individuals in their species. Playbacks can also be used to simulate
intruders in the territory of an individual and can be applied to many other
experimental situations in both the field and laboratory.
The methods used by ethologists are as varied as the problems they in-
vestigate. Because the skill of the observer is still a vital link in the investi-
gation of animal behavior, ethology remains one of the more approachable
areas of scientific investigation.
Uses of Ethology
The investigation of animal behavior has a number of benefits, both practi-
cal and abstract. Some animals are pests, and knowledge of their behavior
can be used to manage them. For example, synthetic pheromones have
been used to attract members of some insect species. The insects may then
be sampled or killed, depending upon the application. To the extent that
researchers develop behaviorally based pest management strategies and
reduce pesticide use, they will be promoting our own safety as well as that
of other species.
It is sometimes important for humans to be able to understand the com-
munication signals of other species and the characteristics of their sensory
perception. The knowledgeable individual can recognize the cues that in-
dicate risk that a dog might bite, for example, and can also avoid behavior
that the dog will regard as threatening. Understanding the behavior of the
wild animals most likely to be encountered in one’s neighborhood is an im-
portant factor in peaceful coexistence.
The study of animal behavior is providing one of the more fascinating
areas of evolutionary biology. Ethology has demonstrated more effectively
220
Ethology
than most fields of study how diverse the solutions to a given problem can
be and has provided insight into human behavior from a biological per-
spective. Knowledge of animal behavior also enriches human lives simply
by satisfying some of our natural curiosity about animals.
See also: Altruism; Communication; Defense mechanisms; Displays; Habit-

uation and sensitization; Herbivores; Hierarchies; Insect societies; Isolating
mechanisms; Mammalian social systems; Migration; Mimicry; Omnivores;
Pheromones; Poisonous animals; Predation; Reproductive strategies; Terri-
toriality and aggression.

Allen, Colin, and Mark Bekoff. Species of Mind: The Philosophy and Biology of
Cognitive Ethology. Cambridge, Mass.: MIT Press, 1997.
Barrows, Edward M. Animal Behavior Desk Reference. 2d ed. Boca Raton,
Fla.: CRC Press, 2000.
Clutton-Brock, T. H., and Paul Harvey, eds. Readings in Sociobiology. San
Francisco: W. H. Freeman, 1978.
Davies, Nicholas B., and John R. Krebs. An Introduction to Behavioral Ecol-
ogy. 4th ed. Boston, Mass.: Blackwell Scientific Publications, 1997.
Goldsmith, Timothy H. The Biological Roots of Human Nature: Forging Links
Between Evolution and Behavior. New York: Oxford University Press,
1991.
Krebs, J. R., and N. B. Davies, eds. Behavioral Ecology: An Evolutionary Ap-
proach. 4th ed. Boston: Blackwell Scientific Publications, 1997.
Lehner, Philip N. Handbook of Ethological Methods. 2d ed. New York: Cam-
Wilson, Edward O. Sociobiology. Cambridge, Mass.: The Belknap Press of
Harvard University Press, 1975.
221
EUTROPHICATION
The overenrichment of water by nutrients, eutrophication causes excessive plant

growth and stagnation, which leads to the death of fish and other aquatic life.
T he word “eutrophic” comes from the Greek eu, which means “well,”
and trophikos, which means “food” or “nutrition.” Eutrophic waters
are well nourished, that is, rich in nutrients; they therefore support abun-
dant life. Eutrophication refers to a condition in aquatic systems (ponds,
lakes, and streams) in which nutrients are so abundant that plants and al-
gae grow uncontrollably and become a problem. The plants die and de-
compose, and the water becomes stagnant. This ultimately causes the
death of other aquatic animals, particularly fish, that cannot tolerate such
conditions. Eutrophication is a major problem in watersheds and water-
ways, such as the Great Lakes and Chesapeake Bay, that are surrounded by
urban populations.
The stagnation that occurs during eutrophication is attributable to the
activity of microorganisms growing on the dead and dying plant material
in water. As they decompose the plant material, microbes consume oxygen
faster than it can be resupplied by the atmosphere. Fish, which need oxygen
in the water to breathe, become starved for oxygen and suffocate. In addi-
tion, noxious gases such as hydrogen sulfide (H2S) can be released during
the decay of the plant material. The hallmark of a eutrophic environment is
one that is plant-filled, littered with dead aquatic life, and smelly.
Eutrophication is actually a natural process that occurs as lakes age and
fill with sediment, as deltas form, and as rivers seek new channels. The
main concern with eutrophication in natural resource conservation is that
human activity can accelerate the process and can cause it to occur in previ-
ously clean but nutrient-poor water. This is sometimes referred to as cul-
tural eutrophication. For example, there is great concern with eutrophica-
tion in Lake Tahoe. Much of Lake Tahoe’s appeal is its crystal-clear water.
Unfortunately, development around Lake Tahoe is causing excess nutri-
ents to flow into the lake and damaging the very thing that attracts people
to the lake.
Roles of Nitrogen and Phosphorus

Nitrogen and phosphorus are the key nutrients involved in eutrophica-
tion, although silicon, calcium, iron, potassium, and manganese can be im-
222
Eutrophication
portant. Nitrogen and phosphorus are essential in plant and animal growth.
Nitrogen compounds are used in the synthesis of amino acids and pro-
teins, whereas phosphate is found in nucleic acids and phospholipids. Ni-
trogen and phosphorus are usually in limited supply in lakes and rivers.
Plants and animals get these nutrients through natural recycling in the
water column and sediments and during seasonal variations, as algae
and animals decompose, fall to the lower depths, and release their nu-
trients to be reused by other organisms in the ecosystem. A limited sup-
ply of nutrients—as well as variations in optimal temperature and light
conditions—prevents any one species of plants or animals from dominat-
ing a water ecosystem.
Although nutrient enrichment can have detrimental effects on a water
system, an increased supply of nitrogen and phosphorus can have an ini-
tial positive effect on water productivity. Much like fertilizers added to a
lawn, nutrients added to lakes, rivers, or oceans stimulate plant and ani-
mal growth in the entire food chain. Phytoplankton—microscopic algae
that grow on the surface of sunlit waters—take up nutrients directly and
are able to proliferate. Through photosynthesis, these primary producers
synthesize organic molecules that are used by other members of the eco-
system. Increased algal growth thus stimulates the growth of zooplank-
ton—microscopic animals that feed on algae and bacteria—as well as
macroinvertebrates, fish, and other animals and plants in the food web. In-
deed, many fisheries have benefitted from lakes and oceans that are pro-
ductive.
Oxygen Depletion
When enough nutrients are added to a lake or river to disrupt the natural
balance of nutrient cycling, however, the excess nutrients effectively become
pollutants. The major problem is that excess nutrients encourage profuse
growth of algae and rooted aquatic weeds, species that can quickly take
advantage of favorable growth conditions at the expense of slower-grow-
ing species. Algae convert carbon dioxide and water into organic mole-
cules during photosynthesis, a process that produces oxygen. When large
blooms of algae and other surface plants die, however, they sink to the bot-
tom of the water to decompose, a process that consumes large amounts of
oxygen. The net effect of increased algae production, therefore, is depletion
of dissolved oxygen in the water, especially during midsummer.
Reduced oxygen levels (called hypoxia) can have dire consequences for
lakes and rivers that support fish and bottom-dwelling animals. Oxygen
depletion is greatest in the deep bottom layers of water, because gases from
the oxygen-rich surface cannot readily mix with the lower layers. During
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Eutrophication
summer and winter, oxygen depletion in eutrophic waters can cause mas-
sive fish kills. In extreme cases of eutrophication, the complete depletion of
oxygen (anoxia) occurs, leading to ecosystem crashes and irreversible
damage to plant and animal life. Oxygen depletion also favors the growth
of anaerobic bacteria, which produce hydrogen sulfide and methane gases,
leading to poor water quality and taste.
Excessive algal and plant growth has other negative effects on a water
system. Algae and plants at the surface block out sunlight to plants and an-
imals at the lower depths. Loss of aquatic plants can affect fish-spawning
areas and encourage soil erosion from shores and banks.
Eutrophication often leads to loss of diversity in a water system, as high
nutrient conditions favor plants and animals that are opportunistic and
short-lived. Native sea grasses and delicate sea plants often are replaced by
hardier weeds and rooted plants. Carp, catfish, and bluegill fish species re-
place more valuable coldwater species such as trout.
Thick algal growth also increases water turbidity and gives lakes and
ponds an unpleasant pea-soup appearance. As algae die and decay, they
wash up on shores in stinking, foamy mats.
Algal blooms of unfavorable species can produce toxins that are harm-
ful to fish, animals, and humans. These toxins can accumulate in shellfish
and have been known to cause death if eaten by humans. So-called red
tides and brown tides are caused by the proliferation of unusual forms of
algae, which give water a reddish or tealike appearance and in some cases
produce harmful chemicals or neurotoxins.
Assessing Eutrophication
While eutrophication effects are generally caused by nutrient enrichment
of a water system, not all cases of nutrient accumulation lead to increased
productivity. Overall productivity is based on other factors in the water
system, such as grazing pressure on phytoplankton, the presence of other
chemicals or pollutants, and the physical features of a body of water. Eu-
trophication occurs mainly in enclosed areas such as estuaries, bays, lakes,
and ponds, where water exchange and mixing are limited. Rivers and
coastal areas with abundant flushing generally show less phytoplankton
growth from nutrient enrichment because their waters run faster and mix
more frequently. On the other hand, activities that stir up nutrient-rich sed-
iments from the bottom, such as development along coastal waters, recre-
ational activities, dredging, and storms, can worsen eutrophication pro-
cesses.
The nutrient status of a lake or water system is often used as a mea-
sure of the extent of eutrophication. For example, lakes are often classified
224
Eutrophication
as oligotrophic (nutrient-poor), eutrophic (nutrient-rich), or mesotrophic

(moderate in nutrients) based on the concentrations of nutrients and the
physical appearance of the lake. Oligotrophic lakes are deep, clear, and un-
productive, with little phytoplankton growth, few aquatic rooted plants,
and high amounts of dissolved oxygen. In contrast, eutrophic lakes are
usually shallow and highly productive, with extensive aquatic plants and
sedimentation. These lakes have high nutrient levels, low amounts of dis-
solved oxygen, and high sediment accumulation on the lake bottom. They
often show sudden blooms of green or blue-green algae (or blue-green bac-
teria, cyanobacteria) and support only warm-water fish species.
Mesotrophic lakes show characteristics in between those of unproduc-
tive oligotrophic waters and highly productive eutrophic waters. Meso-
trophic lakes have moderate nutrient levels and phytoplankton growth
and some sediment accumulation; they support primarily warm-water
fish species. As a lake naturally ages over hundreds of years, it usually (but
not always) gets progressively more eutrophic, as sediments fill in and
eventually convert it to marsh or dryland. Nutrient enrichment from hu-
man sources can speed this process greatly.
Limiting Damage
The negative effects of eutrophication can be reduced by limiting the
amount of nutrients—in most cases nitrogen and phosphorus—from en-
tering a water system. Nutrients can enter water bodies through streams,
rivers, groundwater flow, direct precipitation, and dumping and as partic-
ulate fallout from the atmosphere. While natural processes of eutrophica-
tion are virtually impossible to control, eutrophication from human activ-
ity can be reduced or reversed.
Phosphorus enrichment into water systems occurs primarily as the re-
sult of wastewater drainage into a lake, river, or ocean. Phosphate is com-
mon in industrial and domestic detergents and cleaning agents. Mining
along water systems is also a major source of phosphorus. When phospho-
rus enters a water system, it generally accumulates in the sediments.
Storms and upwelling can stir up sediments, releasing phosphorus. Treat-
ment of wastewater to remove phosphates and the reduction of phos-
phates in detergents have helped to reduce phosphorus enrichment of
water systems.
Nitrogen enrichment is harder to control; it is present in many forms, as
ammonium, nitrates, nitrites, and nitrogen gas. The major sources of nitro-
gen eutrophication are synthetic fertilizers, animal wastes, and agricul-
tural runoff. Some algae species can also fix atmospheric nitrogen directly,
converting it to biologically usable forms of nitrogen. Since the atmosphere
225
Eutrophication
contains about 78 percent nitrogen, this can be a major source of nitrogen

enrichment in waters that already have significant algal populations.
Efforts to control nitrogen and phosphorus levels have examined both
point and nonpoint sources of nutrient loading. Point sources are concen-
trated, identifiable sites of nutrients that include municipal sewage-treat-
ment plants, feed lots, food-processing plants, pulp mills, laundry deter-
gents, and domestic cleaning agents. Nonpoint, or diffuse, sources of
nutrients include surface runoff from rainwater, fertilizer from agricultural
land and lawns, eroded soil, and roadways.
Linda Hart and Mark S. Coyne
See also: Acid deposition; Biomagnification; Erosion and erosion control;

Food chains and webs; Invasive plants; Lakes and limnology; Nutrient cy-
cles; Ocean pollution and oil spills; Pesticides; Phytoplankton; Pollution
effects; Slash-and-burn agriculture; Waste management.

Brönmark, Christer. The Biology of Lakes and Ponds. New York: Oxford Uni-
Cole, Gerald A. Textbook of Limnology. 4th ed. Prospect Heights, Ill.:
Waveland Press, 1994.
Harper, David. Eutrophication of Freshwaters: Principles, Problems, and Resto-
ration. London: Chapman & Hall, 1992.
Hinga, Kenneth, Heeseon Jeon, and Noelle F. Lewis. Marine Eutrophication
Review. Silver Springs, Md.: U.S. Department of Commerce, National
Oceanic and Atmospheric Administration, Coastal Ocean Office, 1995.
Horne, Alexander J., and Charles R. Goldman. Limnology. 2d ed. New York:
McGraw-Hill, 1994.
Schramm, Winfrid, and Pieter H. Nienhuis, eds. Marine Benthic Vegetation:
Recent Changes and the Effects of Eutrophication. Berlin: Springer, 1996.
Wetzel, Robert G. Limnological Analyses. 3d ed. New York: Springer, 2000.
226
EVOLUTION:
DEFINITION AND THEORIES
Types of ecology: Evolutionary ecology; Paleoecology; Speciation
Evolution is change in species through time. From a one-celled ancestor, many bil-
lions of species have evolved into a grand diversity of life-forms that populate earth
and interact in ways that will affect the evolution of future life-forms.
Inheritance and Natural Selection

“Evolution” comes from the Latin word meaning “to unroll.” In a general
sense, it refers to any change through time, but it is often restricted to bio-
logical change. For most of human history, the universe was thought to be
unchanging. Then, in the eighteenth century, expeditions to new continents
and the discovery of extinct fossil animals convinced many people that the
biological world was not as unchanging as had been thought. There could
be no proof for this hypothesis, however, until an explanation of how such
change occurred could be found. In 1809, Jean-Baptiste Lamarck became
the first to propose an explanation; his theory was based on the inheritance
of acquired traits. According to this theory, giraffes, for example, obtained
long necks because individual giraffes stretched their neck muscles more
and more to reach ever higher leaves, and the longer necks were passed on
to the offspring. This idea was quickly shown to be false by experiment.
Traits acquired during an individual’s lifetime (such as larger muscles ac-
quired through weightlifting) are not passed on to offspring.
It was 1859 when Charles Darwin proposed what is now known to be
the actual process by which evolution occurs: natural selection. This pro-
cess can be divided into three steps: Individuals in a species vary in their
traits; some individuals will have more offspring than others, depending
on how advantageous their particular traits are; advantageous traits will
increase in a species, and disadvantageous traits will be lost through time
and as the environment changes. For example, climatic change may cause
a forested environment to become a snowy tundra. Creatures with dark
coats are best off in a forest because of the concealment of the dark, shad-
owy environment, but as the lighter, snowy environment becomes domi-
nant, lighter individuals will have an advantage because they are more
easily concealed. Over a long period, enough changes will accumulate in a
group that an observer might say that a new species had been created. This
process has often been called “survival of the fittest,” but the “fittest” or-
227
Evolution: definition and theories
ganisms are not always the fastest, fiercest, or even most competitive. For
example, animals that cooperate with other animals or are the most timid
and conceal themselves readily may survive more often and produce more
offspring.
When the whole species changes at once, “nonbranching” evolution oc-
curs. Many species, however, have wide ranges and occur in many differ-
ent geographic areas, so often only some populations of a species are sub-
jected to environmental changes. That is an important point because it
explains how so many species can be created from one ancestral species.
“Branching” evolution is especially common when one of the populations
becomes cut off from the others by a barrier of some kind. For example, a
new river may form. This river prevents interbreeding and allows each
population to form its own pool of traits. In time, differences between the
two environments will cause the two populations to become so distinct
that they form two different species.
Species have hitherto been described as groups that are visibly distinct
enough to be distinguishable from one another; however, there is a much
more objective definition of species, based on the criterion of interbreed-
ing. To a biologist, members of a species can produce fertile offspring only
when they breed with other members. Therefore, a new species has
evolved not when it “looks” sufficiently different from its ancestors or
neighboring populations but when it can no longer successfully interbreed
with them. This definition of reproductive isolation is important because
many closely related species look quite similar yet cannot successfully in-
terbreed.
Evolution by natural selection explains not only how species have
changed but also why they are so well adapted to their surroundings: The
best-adapted individuals have the most offspring. Further, it explains
some crucial aspects of basic anatomy, such as why vestigial, or “remnant,”
organs exist: They are in the process of being lost. For example, the now-
useless human appendix was once an important part of the human diges-
tive tract. Also, it explains why many organisms have similar organs that
are used for different purposes, such as five-fingered hands on humans
and five-digit organs on bat wings. Such “homologous” organs have been
modified from a common ancestor. This modification also explains why
many organisms pass through similar embryonic stages; human embryos,
for example, have tails and gills like a fish.
Laws of Heredity
After Darwin proposed natural selection as the process of evolution, it was
readily accepted, and most scientists have accepted it ever since. The ex-
228
planation was incomplete, however, in one major area: Darwin could not
explain how variation was produced or passed on. The laws of heredity
were discovered by Gregor Mendel in 1866 while Darwin was wrestling
with this problem. Mendel’s work lay unnoticed until the early twentieth
century, when other scientists independently discovered the gene as the
basic unit of heredity. Genes are now known to be molecular “blueprints”
that are repeatedly copied within each cell. They contain instructions on
how to build the organism and how to maintain it.
Genes are passed on to the offspring when a sperm and an egg cell
unite. The resulting fertilized egg consists of one cell that contains all the
genes on strands, or chromosomes, in the cell nucleus. The chromosomes
occur in pairs such that one member of each pair is from the father and one
is from the mother. As growth occurs, certain genes in each cell will be bio-
chemically “read” and will give instructions on what happens next. Genes
are composed of the molecule deoxyribonucleic acid (DNA), which is
shaped like a twisted ladder and is copied when the “ladder” splits in half
at the middle of the “rungs.” Once the instructions are copied, they are car-
ried outside the cell nucleus by messenger molecules, which proceed to
build proteins (such as enzymes and muscle tissue) using the rungs as a
blueprint.
With this added knowledge of genes as the units of heredity, evolution-
ists could see that natural selection acting on individuals selects not only
traits but also the genes that serve as blueprints for those traits. Therefore,
as well as being a change in a species’ traits through time, evolution is also
often defined as a change in the “gene pool” of a species. The gene pool is
the total of all the genes contained in a species. Individual variation in a
gene pool originally arises via mutations, errors made in the DNA copying
process. Usually, mutation involves a change in the DNA sequence that
causes a change in the genetic instructions.
Most mutations have little effect, which is fortunate because those that
are expressed generally kill or handicap the offspring. That occurs since
any organism is a highly integrated, complex system, and any major alter-
ations to it are therefore likely to disrupt it. Nevertheless, rare improve-
ments do occur, and it is these that are passed on and become part of the
breeding gene pool. Although mutations provide the ultimate source of
variation, the sexual recombination of genes provides the more immediate
source. Each organism has a unique combination of genes, and it is the fit-
ness of this combination that determines how well those genes survive and
are passed on. Although brothers and sisters have the same parents, they
are not alike because genes are constantly shuffled and reshuffled in the
production of each sperm and egg cell.
229
The Geologic Time Scale
MYA Eon Era Period Epoch Developments

Quaternary Holocene Ice Age ends. Human activities begin to
0.01 (1.8 mya-today) (11,000 ya-today) impact biosphere.
Pleistocene Ice Age in northwest Europe, Siberia,
(1.8 mya-11,000 ya) and North America. Plants migrate in
response. New speciations lead to
1.8 modern plants. Modern humans evolve.
Tertiary Pliocene Cooling period leads to Ice Age.
(65-1.8 mya) (5-1.8 mya)
(65 mya-today)
5
Cenozoic
Miocene Erect-walking human ancestors

23 (23-5 mya)
Oligocene Primate ancestors of humans
38 (38-23 mya)
Eocene Intense mountain building occurs (Alps,
(54-38 mya) Himalaya, rockies); modern mammals
appear (rodents, hoofed animals);
54 diverse conferous forests
Paleocene Cretaceous-Tertiary even leads to
(65-54 mya) dinosarus’ extinction c. 65 mya; seed
ferns; bennettites and caytonias dief off;
65 ginkgoes decline; decidous plants rise.
Phanerozoic Eon
(544 mya-today)
Cretaceous (146-65 mya) Breakup of supercontinents into tosay’s

form; birds appear. Cycads, other
gymnosperms still widespread, but
angiosperms dominate by 90 mya;
animal-aided pollination begins to
146 evolve.
(245-208 mya)
Jurassic (208-146 mya) Earliest mammals; ginkgoes thrive in

Mesozoic
moister areas; drier climates in North

and South America, parts of Africa,
central Asia; rise of modern
gymnosperms, such as junipers, pine
trees. Earliest angiosperm fossil (from
208 China) dates from end of this period.
Triassic (245-208 mya) Diminished land vegetation, with lack
of variation reflecting global frost-free
climate; gymnosperms dominate,
bennettites and gnetophytes appear;
245 dinosaurs develop.
Permian (286-245 mya) Permian extinction even initiates drier,
colder period; supercontinent Pangaea
has formed; tree-sized lycophytes, club
(544-245 mya)
mosses, cordaites die off; horsetails,

Paleozoic
286 peltasperms, cycads, conifers dominate.

Carboniferous Pennsylvanian Gymnosperms, ferns, calamites,
325 (360-286 mya) (325-286 mya) lycopods thrive (first seed plants);
reptiles appear. Plant life diverse, from
Mississippian
small creeping forms to tall forest trees.
(360-325 mya)
coal beds form in swamp forests from
360 the dominant seedless vascular plants.
MYA Eon Era Period Epoch Developments

410 Devonian (410-360 mya) Club mosses, early ferns,
lycophytes, progymnosperms;
amphibians, diverse insects;
horsetails, gymnosperms present
by end of period.
Phanerozoic Eon
(244 mya-today)
440 Silurian (440-410 mya) Early land plants: nonvascular
(544-245 mya)
bryophytes (mosses, hornworts),
Paleozoic
followed by seedless vascular
plants in now-extinct phyla
Rhyniophyta, Zosterophyllophyta,
Trimerophytophyta.
505 Ordovician (505-440 mya) Life colonizes land; earliest
vertebrates appear in fossil record.
544 Cambrian Tommotian Cambrian diversification of life
(544-505 mya) (530-527 mya)
900 Neoproterozoic Vendian Age of algae, earliest invertebrates
(2,500-544 mya)
(900-544 mya) (650-544 mya)

Precambrian Time
Proterozoic
(4,500-544 mya)
1600 Mesoproterozoic (1,600-900 mya) Eukaryotic life established.

2500 Paleoproterozoic (2,500-1,600 mya) Transition from prokaryotic to
eukaryotic life leads to
multicellular organisms
3800 Archaean (3,800-2,500 mya) Microbial life (anaerobic and
cyanobacteria) as early as 3.5 bya.
4500 Hadean (4,500-3,800 mya) Earth forms 4.5 bya.
Source: Data on time periods in this version of the geologic time scale are based on new
findings in the last decade of the twentieth century as presented by the Geologic Society
of America, which notably moves the transition between the Precambrian and Cambrian
times from 570 mya to 544 mya.
Notes: bya=billions of years ago; mya = millions of years ago; ya = years ago.
Rates and Patterns of Evolution

A major area of debate is how fast evolution occurs. Some scientists believe
that most evolution occurs rapidly. This view has been called “punctuated”
evolution. Another group argues that evolution is more often gradual, as
Darwin originally proposed. To some extent, this disagreement is a matter of
different perspectives. A geneticist working with flies in a laboratory would
see the evolution of a new species in ten thousand years as very slow. To a
paleontologist, however, who often deals with fossil species lasting millions
of years, ten thousand years is brief indeed. Nevertheless, there is more to
the debate than perspective alone. Punctuationists argue not only that evo-
lution is rapid but also that species have such tightly integrated gene pools
that virtually no change at all occurs during most of a species’ existence.
In contrast, gradualists view species as being much less integrated, so
that change can be a continuous process. The fossil record at first glance
231
seems to support the punctuated view. The majority of species show very
little change for long spans of time and then either disappear or rapidly
give rise to another species. The fossil record is very incomplete, however,
being full of gaps where no fossils were deposited. As a result, it is often
impossible to tell whether the “rapid” change in species is real or only fol-
lows a gap in what was actually a gradual sequence. Also, fossils represent
only part of the original organism—usually only the hard parts, such as
shells, bones, or teeth. Therefore, any changes in soft anatomy, such as tis-
sues or biochemistry, are lost, making it impossible to say with certainty
that no change occurred. Whatever the outcome of the debate, all scientists
agree that evolutionary rates vary.
In addition to the rate of evolution, much has also been written about
the patterns produced by evolution since life arose about 3.5 million years
ago. Evolutionary trends are directional changes seen in a group. The most
common trend, found in many groups, is an increase in size. Another
trend, seen mainly in mammals, is an increase in brain size. Life as a whole
has shown an increase in total diversity and complexity. These trends,
however, are only statistical tendencies. They are not inevitable “laws,” as
many have misinterpreted them in the past. Often, groups do not show
them, and in those that do, the change is not constant and may reverse it-
self at times. Finally, trends are often interrupted by mass extinctions. At
least five times in the past 600 million years, more than 50 percent of all the
species on the earth have been wiped out by catastrophes of different
kinds, from temperature changes to impacts of huge meteorites.
Study of Fossils
Fossils, the remains of former life, provide the only record of most evolu-
tion, because more than 99 percent of all species that have ever existed are
now extinct. Paleontology is the study of fossils. Such study begins with
identification of the remains—usually hard parts, such as bones—and
ends with measurement of fossil size, shape, and abundance. The extreme
incompleteness of the fossil record is a major obstacle to this method, since
only some parts are preserved, and these are from strictly limited periods
of the evolutionary past. Nevertheless, many evolutionary lineages can be
traced through time. Indeed, refined measurements of rate and direction of
anatomical change are often possible when used in conjunction with dat-
ing techniques.
The study of living organisms permits observation of the complete or-
ganism. Comparative anatomy reveals similarities among related species
and shows how evolution has modified them since they separated from
their common ancestor. For example, humans and chimpanzees are ex-
232
tremely similar in their organ and muscle anatomy. This method is not lim-
ited to comparison of adults but includes earlier stages of development as
well. Comparative embryology often shows anatomical similarities, such
as those between humans and other vertebrates. For example, the human
embryo goes through a stage with gills and a tail, resembling stages of an
amphibian embryo. Comparative biochemistry is also very useful, reveal-
ing similarities in proteins and many other molecules. Such comparisons
are based on differences in molecular sequences, such as amino acids. Mo-
lecular “clocks” are sometimes calculated in this manner. More distantly
related species are thought to have more differences. The accuracy of such
clocks, however, is hotly debated.
A major technique is DNA sequencing, whereby the exact genetic infor-
mation is read directly from the gene. This method will greatly add to
knowledge of evolutionary relationships, although it is expensive and
time-consuming. Biogeography, or the distribution of organisms in nature,
is a method that Darwin used and that is still important today. This tech-
nique often reveals populations (races) within a species’ overall range that
differ from one another because they inhabit slightly different geographic
areas. These populations give the scientist a “snapshot” of evolution in
progress. Given more time, many of these races would eventually become
different species.
Artificial breeding is a method of directly manipulating evolution. The
most widely used experimental organism for this purpose is the fruit fly,
which is used in part because of its exceptionally large chromosomes; they
make the genes easy to identify. A common experiment is to subject the
flies to radiation or chemicals that cause mutations and then to analyze the
effects. The gene pool is then subjected to extreme artificial selection as the
experimenter allows only certain individuals to breed. For example, only
those with a gene for a certain kind of wing may reproduce. Although such
experiments have often altered the organisms’ gene pools and created new
varieties within the species, no truly new species has ever been created in
the laboratory. Apparently, more time is needed to produce a new species.
Outside the laboratory, artificial breeding has been done for thousands of
years. Food plants and domesticated animals have had much of their evo-
lution controlled by humans. Analysis of the effect of this breeding on the
organisms’ gene pools is the most complete and direct method of studying
evolution.
Significance
The study of fossils has been a major tool in understanding Earth’s history.
This understanding has allowed more efficient exploitation of the earth’s
233
resources. For example, petroleum and coal provide the major energy re-
sources today and were both formed by organisms of the past. Petroleum
comes from the biochemicals of marine organisms, and coal comes from
fossilized plants. Most paleontologists are employed in the costly search
for these “fossil fuels,” and knowing the evolutionary history of these
groups helps to determine the most productive places to search. Fossils
also form nonenergy resources. Limestone is used in many processes, from
making cement to making steel. Most limestone is composed of the fossil-
ized remains of seashells and other marine skeletons. Phosphate minerals,
essential for fertilizers in almost all forms of agriculture, come from marine
fossil deposits as well.
Darwin’s theory of natural selection caused a violent reaction through-
out much of the world when it was applied in social contexts (to which
Darwin himself disagreed) as “social Darwinism.” The notion that humans
evolved from lower life-forms such as the ape was truly revolutionary. In-
stead of creatures of a divine plan, humans were now seen as products of
natural, sometimes “random,” processes. The impact of this realization on
ethics, the arts, and society in general is still being felt. Evolution, however,
does not necessarily conflict with religion, as is often thought. Science
seeks to find out only how things happen, not the ultimate reasons why
they happen. Therefore, most major religions have reconciled their tenets
with the fact of evolution by viewing natural selection as simply a mecha-
nism employed by God to meet his ends.
Michael L. McKinney

lution; Colonization of the land; Convergence and divergence; Dendro-
chronology; Development and ecological strategies; Evolution: history;
Evolution of plants and climates; Extinctions and evolutionary explosions;
Gene flow; Genetic drift; Genetically modified foods; Isolating mecha-
Paleoecology; Population genetics; Punctuated equilibrium vs. gradual-
ism; Speciation; Species loss.

Dawkins, Richard. The Blind Watchmaker: Why the Evidence of Evolution Re-
veals a Universe Without Design. New York: W. W. Norton, 1988.
Futuyma, Douglas J. Evolutionary Biology. Sunderland, Mass.: Sinauer As-
sociates, 1986.
McNamara, K. J. Shapes of Time: The Evolution of Growth and Development.
Baltimore: Johns Hopkins University Press, 1997.
234
Schopf, J. William, ed. Major Events in the History of Life. Boston: Jones and
Bartlett, 1992.
Stanley, Steven M. The New Evolutionary Timetable: Fossils, Genes, and the Or-
igin of Species. New York: Basic Books, 1981.
Stebbins, G. Ledyard. Darwin to DNA, Molecules to Humanity. New York:
W. H. Freeman, 1982.
235
EVOLUTION: HISTORY
Types of ecology: Evolutionary ecology; History of ecology
Evolution is the theory that biological species undergo sufficient change with time
to give rise to new species. The development of the theory of evolution has contrib-
uted much to two later scientific disciplines, genetics and ecology, by providing ex-
planations for the adaptations and interrelationships of species from early times to
the present.
T he concept of evolution has ancient roots. Anaximander suggested in

the sixth century b.c.e. that life had originated in the seas and that
humans had evolved from fish. Empedocles (c. 450 b.c.e.) and Lucretius
(c. 96-55 b.c.e.), in a sense, grasped the concepts of adaptation and natural
selection. They taught that bodies had originally formed from the random
combination of parts, but that only harmoniously functioning combina-
tions could survive and reproduce. Lucretius even said that the mythical
centaur, half horse and half human, could never have existed because the
human teeth and stomach would be incapable of chewing and digesting
the kind of grassy food needed to nourish the horse’s body.
Early Biological Theory

For two thousand years, however, evolution was considered an impossi-
bility. The theory of forms (also called his theory of ideas) proposed by
Plato (c. 428-348 b.c.e.) gave rise to the notion that each species had an un-
changing “essence” incapable of evolutionary change. As a result, most
scientists from Aristotle (384-322 b.c.e.) to Carolus Linnaeus (1707-1778)
insisted upon the immutability of species.
Many of these scientists tried to arrange all species in a single linear se-
quence known as the scale of being (also called the great chain of being or
scala naturae), a concept supported well into the nineteenth century by
many philosophers and theologians. The sequence in this scale of be-
ing was usually interpreted as a static “ladder of perfection” in God’s cre-
ation, arranged from higher to lower forms. The scale had to be continu-
ous, for any gap would detract from the perfection of God’s creation. Much
exploration was devoted to searching for missing links in the chain, but
it was generally agreed that the entire system was static and incapable
of evolutionary change. Pierre-Louis Moreau de Maupertuis and Jean-
Baptiste Lamarck (1744-1829) were among the scientists who tried to rein-
terpret the scale of being as an evolutionary sequence, but this single-
236
Evolution: history
sequence idea was later replaced by the concept of branching evolution

proposed by Charles Darwin (1809-1882). Georges Cuvier (1769-1832) fi-
nally showed that the major groups of animals had such strikingly differ-
ent anatomical structures that no possible scale of being could connect
them all; the idea of a scale of being lost most of its scientific support as a
result.
The theory that new biological species could arise from changes in exist-
ing species was not readily accepted at first. Linnaeus and other classi-
cal biologists emphasized the immutability of species under the Platonic-
Aristotelian concept of essentialism. Those who believed in the concept of
evolution realized that no such idea could gain acceptance until a suitable
mechanism of evolution could be found. Many possible mechanisms were
therefore proposed. Étienne Geoffroy Saint-Hilaire (1805-1861) proposed
that the environment directly induced physiological changes, which he
thought would be inherited, a theory now known as Geoffroyism. Lamarck
proposed that there was an overall linear ascent of the scale of being but
that organisms could also adapt to local environments by voluntary exer-
cise, which would strengthen the organs used; unused organs would dete-
riorate. He thought that the characteristics acquired by use and disuse
would be passed on to later generations, but the inheritance of acquired
characteristics was later disproved. Central to both these explanations was
the concept of adaptation, or the possession by organisms of characteristics
that suit them to their environments or to their ways of life. In eighteenth
century England, the Reverend William Paley (1743-1805) and his numer-
ous scientific supporters believed that such adaptations could be explained
only by the action of an omnipotent, benevolent God. In criticizing Lamarck,
the supporters of Paley pointed out that birds migrated toward warmer cli-
mates before winter set in and that the heart of the human fetus had fea-
tures that anticipated the changes of function that take place at birth. No
amount of use and disuse could explain these cases of anticipation, they
claimed; only an omniscient God who could foretell future events could
have designed things with their future utility in mind.
Darwin’s Theory
The nineteenth century witnessed a number of books asserting that living
species had evolved from earlier ones. Before 1859, these works were often
more geological than biological in content. Most successful among them
was the anonymously published Vestiges of the Natural History of Creation
(1844), written by Robert Chambers (1802-1871). Books of this genre sold
well but contained many flaws. They proposed no mechanism to account
for evolutionary change. They supported the outmoded concept of a scale
237
Evolution: history
of being, often as a single sequence of evolutionary “progress.” In geology,

they supported the outmoded theory of catastrophism, an idea that the his-
tory of the earth had been characterized by great cataclysmic upheavals.
From 1830 on, however, that theory was being replaced by the modern the-
ory of uniformitarianism, championed by Charles Lyell (1797-1875). Dar-
win read these books and knew their faults, especially their lack of a mech-
anism that was compatible with Lyell’s geology. In his own work, Darwin
carefully tried to avoid the shortcomings of these theories. Eventually, he
brought about the greatest revolution in biological thought by proposing
both a theory of branching evolution and a mechanism of natural selection
to explain how it occurred.
Much of Darwin’s evidence was gathered during his voyage around the
world aboard HMS Beagle between 1831 and 1836. Darwin’s stop in the
Galápagos Islands and his study of tortoises and finchlike birds on these is-
lands is usually credited with convincing him that evolution was a branch-
ing process and that adaptation to local environments was an essential part
of the evolutionary process. Adaptation, he later concluded, came about
through natural selection, a process that led to the deaths of maladapted
variations and allowed only the well-adapted ones to survive and pass on
their hereditary traits. After returning to England from his voyage, Darwin
raised pigeons, consulted with various animal breeders about changes in
domestic breeds, and investigated other phenomena that later enabled
him to demonstrate natural selection and its power to produce evolution-
ary change.
Darwin delayed the publication of his book for seventeen years after he
wrote his first manuscript version. He might have waited even longer, ex-
cept that his hand was forced. From the East Indies, another British scien-
tist, Alfred Russel Wallace (1823-1913), had written a description of an
identical theory and submitted it to Darwin for his comments. Darwin
showed Wallace’s letter to Lyell, who urged that both Darwin’s and
Wallace’s contributions be published, along with documented evidence
showing that both had arrived at the same ideas independently. Darwin’s
great book, On the Origin of Species by Means of Natural Selection, was pub-
lished in 1859, and it quickly won most of the scientific community’s sup-
port of the concept of branching evolution. In his later years, Darwin also
published The Descent of Man and Selection in Relation to Sex (1871), in which
he outlined his theory of sexual selection. According to this theory, the
agent that determines the composition of the next generation may often be
the opposite sex. An organism may be well adapted to live, but unless it
can mate and leave offspring, it will not contribute to the next or to future
generations.
238
Evolution: history
After Darwin
In the early 1900’s, the rise of Mendelian genetics (named for botanist
Gregor Mendel, 1822-1884) initially resulted in challenges to Darwinism.
Hugo de Vries (1848-1935) proposed that evolution occurred by random
mutations, which were not necessarily adaptive. This idea was subse-
quently rejected, and Mendelian genetics was reconciled with Darwinism
during the period from 1930 to 1942. According to this modern synthetic
theory of evolution, mutations initially occur at random, but natural selec-
tion eliminates most of them and alters the proportions among those that
survive. Over many generations, the accumulation of heritable traits pro-
duces the kind of adaptive change that Darwin and others had described.
The process of branching evolution through speciation is also an important
part of the modern synthesis.
The branching of the evolutionary tree has resulted in the proliferation
of species from the common ancestor of each group, a process called adap-
tive radiation. Ultimately, all species are believed to have descended from
a single common ancestor. Because of the branching nature of the evolu-
tionary process, no one evolutionary sequence can be singled out as repre-
senting any overall trend; rather, there have been different trends in differ-
ent groups. Evolution is also an opportunistic process, in the sense that it
follows the path of least resistance in each case. Instead of moving in
straight lines toward a predetermined goal, evolving lineages often trace
meandering or circuitous paths in which each change represents a momen-
tary increase in adaptation. Species that cannot adapt to changing condi-
tions die out and become extinct.
Evolutionary biology is itself the context into which all the other biolog-
ical sciences fit. Other biologists, including physiologists and molecular bi-
ologists, study how certain processes work, but it is evolutionists who
study the reasons that these processes came to work in one way and not an-
other. Organisms and their cells are built one way and not another be-
cause their structures have evolved in a particular direction and can only
be explained as the result of an evolutionary process. Not only does each
biological system need to function properly, but it also must have been
able to achieve its present method of functioning as the result of a long, his-
torical, evolutionary process in which a previous method of functioning
changed into the present one. If there were two or more ways of accom-
plishing the same result, a particular species used one of them because its
ancestors were more easily capable of evolving this one method rather
than another.
Eli C. Minkoff
239
Evolution: history

chronology; Development and ecological strategies; Ecology: history; Eco-
systems: definition and history; Evolution: definition and theories;
Evolution of plants and climates; Extinctions and evolutionary explosions;
Gene flow; Genetic drift; Genetically modified foods; Isolating mecha-
Paleoecology; Population genetics; Punctuated equilibrium vs. gradual-
ism; Speciation; Species loss.

Bowler, Peter J. Evolution: The History of an Idea. Rev. ed. Berkeley: Univer-
sity of California Press, 1989.
_______. Life’s Splendid Drama: Evolutionary Biology and the Reconstruction of
Life’s Ancestry, 1860-1940. Chicago: University of Chicago Press, 1996.
Brandon, Robert N. Concepts and Methods in Evolutionary Biology. New
Grant, Verne. The Evolutionary Process: A Critical Study of Evolutionary The-
ory. 2d ed. New York: Columbia University Press, 1991.
Minkoff, Eli C. Evolutionary Biology. Reading, Mass.: Addison-Wesley,
1983.
Zimmer, Carl. Evolution: The Triumph of an Idea. New York: HarperCollins,
2001.
240
EVOLUTION OF PLANTS
AND CLIMATES
Types of ecology: Evolutionary ecology; Paleoecology; Speciation
As a result of prehistoric events such as the Permian-Triassic extinction event and

the Cretaceous-Tertiary mass extinction event, many plant families and some an-
cestors of extant plant were extinct before the beginning of recorded history.
T he general trend of earth’s plant diversification involves four major

plant groups that rose to dominance from about the Middle Silurian
period to present time. The first major group providing land vegetation
comprised the seedless vascular plants, represented by the phyla
Rhyniophyta, Zosterophyllophyta, and Trimerophytophyta. The second major
group appearing in the late Devonian period was made up of the ferns
(Pterophyta). The third group, the seed plants (sometimes called the Coal
Age plants), appeared at least 380 million years ago (mya). This third
group includes the gymnosperms (Gymnospermophyta), which dominated
land flora for most of the Mesozoic era until 100 mya. The last group, the
flowering angiosperms (Anthophyta), appeared in the fossil record 130
mya. The fossil record also shows that this group of plants was abundant in
most parts of the world within 30 million to 40 million years. Thus, the an-
giosperms have dominated land vegetation for close to 100 million years.
The Paleozoic Era

The Proterozoic and Archean eons have restricted fossil records and pre-
date the appearance of land plants. Seedless, vascular land plants ap-
peared in the middle of the Silurian period (437-407 mya) and are repre-
sented by the rhyniophytes or rhyniophytoids and possibly the Lycophyta
(lycophytes or club mosses). From the primitive rhyniophytes and lyco-
phytes, land vegetation rapidly diversified during the Devonian period
(407-360 mya). Pre-fern ancestors and maybe true ferns (Pterophyta) were
developed by the mid-Devonian. By the Late Devonian the horsetails
(Sphenophyta) and gymnosperms (Gymnospermophyta) were present. By the
end of the period, all major divisions of vascular plants had appeared ex-
cept the angiosperms.
Development of vascular plant structures throughout the Devonian al-
lowed for greater geographical diversity of plants. One such structure was
flattened, planated leaves, which increased photosynthetic efficiency. An-
241
Evolution of plants and climates
other was the development of secondary wood, allowing plants to increase

significantly in structure and size, thus resulting in trees and probably for-
ests. A gradual process was the reproductive development of the seed; the
earliest structures are found in Upper Devonian deposits.
Ancestors of the conifers and cycads appeared in the Carboniferous pe-
riod (360-287 mya), but their documentation is poor in the fossil record.
During the early Carboniferous in the high and middle latitudes, vegeta-
tion shows a dominance of club mosses and progymnosperms (Progymno-
spermophyta). In the lower latitudes of North America and Europe, a greater
diversity of club mosses and progymnosperms are found, along with a
greater diversity of vegetation. Seed ferns (lagenostomaleans, calamo-
pityaleans) are present, along with true ferns and horsetails (Archaeocal-
amites).
Late Carboniferous vegetation in the high latitudes was greatly affected
by the start of the Permo-Carboniferous Ice Age. In the northern middle
latitudes, the fossil record reveals a dominance of horsetails and primitive
seed ferns (pteridosperms) but few other plants.
In northern low latitudes, landmasses of North America, Europe, and
China were covered by shallow seas or swamps and, because they were
close to the equator, experienced tropical to subtropical climatic condi-
tions. The first tropical rain forests appeared there, known as the Coal Mea-
sure Forests or the Age of Coal. Vast amounts of peat were laid down as a
result of favorable conditions of year-round growth and the giant club
mosses’ adaptation to the wetland tropical environments. In drier areas
surrounding the lowlands, forests of horsetails (calamites, sphenophylls),
seed ferns (medullosans, callistophytes, lagenostomaleans), cordaites, and
diverse ferns (including marattialean tree ferns) existed in great abun-
dance.
The Permian period (287-250 mya) marks a major transition of the coni-
fers, cycads, glossopterids, gigantopterids, and the peltasperms from a
poor fossil record in the Carboniferous to significantly abundant land
vegetation. The two most prevalent plant assemblages of the Permian were
the horsetails, peltasperms, cycadophytes, and conifers. The second most
prevalent were the gigantopterids, peltasperms, and conifers. These two
plant assemblages are considered the typical paleo-equatorial lowland
vegetation of the Permian. Other plants, such as the tree ferns and giant
club mosses, were present in the Permian but not abundant. As a result of
the Permian-Triassic extinction event, tropical swamp forests disappeared,
with the extinction of the club mosses; the cordaites and glossopterids dis-
appeared from higher latitudes; and 96 percent of all plant and animal spe-
cies became extinct.
242
The Mesozoic Era

At the beginning of the Triassic period (248-208 mya), a meager fossil re-
cord reveals diminished land vegetation (that is, no coal formed). By the
middle to late Triassic, the modern family of ferns, conifers, and a now-ex-
tinct group of plants, the bennettites (cycadeoids), inhabited most land
surfaces. After the mass extinction, the bennettites moved into vacant low-
land niches. They may be significant because of the similarity of their re-
productive organs to the reproductive organs of the angiosperms.
Late Triassic flora in the equatorial latitudes are represented by a wide
range of ferns, horsetails, pteriosperms, cycads, bennettites, leptostro-
baleans, ginkgos, and conifers. The plant assemblages in the middle lati-
tudes are similar but not as species-rich. This lack of plant variation in low
and middle latitudes reflects a global frost-free climate.
In the Jurassic period (208-144 mya), land vegetation similar to modern
vegetation began to appear, and the ferns of this age can be assigned to
modern families: Dipteridaceae, Matoniaceae, Gleicheniaceae, and Cyatheaceae.
Conifers of this age can also be assigned to modern families: Podocarpaceae,
Araucariaceae (Norfolk pines), Pinaceae (pines), and Taxaceae (yews). These
conifers created substantial coal deposits in the Mesozoic.
During the Early to Middle Jurassic, diverse vegetation grew in the
equatorial latitudes of western North America, Europe, Central Asia, and
the Far East and comprised the horsetails, pteridosperms, cycads, ben-
nettites, leptostrobaleans, ginkgos, ferns, and conifers. Warm, moist condi-
tions also existed in the northern middle latitudes (Siberia and northwest
Canada), supporting Ginkgoalean forests and leptostrobaleans. Desert
conditions existed in central and eastern North America and North Africa,
and the presence of bennettites, cycads, peltasperms, and cheirolepidiacean
conifers there are plant indicators of drier conditions. The southern lati-
tudes had vegetation similar to that of the equatorial latitudes, but owing
to drier conditions, cheirolepidiacean conifers were abundant, ginkgos
scarce. This southern vegetation spread into very high latitudes, including
Antarctica, because of the lack of polar ice.
In the Cretaceous period (144-66.4 mya), arid, subdesert conditions ex-
isted in South America, Central and North Africa, and central Asia. Thus,
the land vegetation was dominated by cheirolepidiacean conifers and
matoniacean ferns. The northern middle latitudes of Europe and North
America had a more diverse vegetation comprising bennettites, cycads,
ferns, peltasperms, and cheirolepidiacean conifers with the southern mid-
dle latitudes dominated by bennettites and cheirolepidiaceans.
A major change in land vegetation took place in the late Cretaceous with
the appearance and proliferation of flowering seed plants, the angio-
243
sperms. The presence of the angiosperms marked the end of the typical
gymnosperm-dominated Mesozoic flora and a definite decline in the
leptostrobaleans, bennettites, ginkgos, and cycads.
During the late Cretaceous in South America, central Africa, and India,
arid conditions prevailed, resulting in tropical vegetation dominated by
palms. The southern middle latitudes were also affected by desert condi-
tions, and the plants that fringed these desert areas were horsetails, ferns,
conifers (araucarias, podocarps), and angiosperms, specifically Notho-
fagus (southern beech). The high-latitude areas were devoid of polar ice;
owing to the warmer conditions, angiosperms were able to thrive. The
most diverse flora was found in North America, with the presence of ever-
greens, angiosperms, and conifers, especially the redwood, Sequoia.
The Cretaceous-Tertiary (K/T) mass extinction event occurred at about
66.4 mya. This event has been hypothesized to be a meteoritic impact;
whatever the cause, at this time an event took place that suddenly induced
global climatic change and initiated the extinction of many species, nota-
bly the dinosaurs. The K/T had a greater effect on plants with many fami-
lies than it did on plants with very few families. Those that did become ex-
tinct, such as the bennettites and caytonias, had been in decline. The
greatest shock to land vegetation occurred in the middle latitudes of North
America. The pollen and spore record just above the K/T boundary in the
fossil record shows a dominance of ferns and evergreens. Subsequent plant
colonization in North America shows a dominance of deciduous plants.
The Cenozoic Era

Increased rainfall at the beginning of the Paleogene-Neogene period (66.4-
1.8 mya) supported the widespread development of rain forests in south-
erly areas. Rain forests are documented by larger leaf size and drip tips at
leaf edge, typical characteristics of modern rain-forest floras.
Notable in this period was the polar Arcto-Tertiary forest flora found in
northwest Canada at paleolatitudes of 75-80 degrees north. Mild, moist
summers alternated with continuous winter darkness, with temperatures
ranging from 0 to 25 degrees Celsius. These climatic conditions supported
deciduous vegetation that included Platanaceae (sycamore), Judlandaceae
(walnut), Betulaceae (birch), Menispermaceae, Cercidophyllaceae, Ulmaceae
(elm), Fagaceae (beech), Magnoliaceae; and gymnosperms such as Taxo-
diaceae (redwood), Cypressaceae (cypress), Pinaceae (pine), and Ginkgoaceae
(gingko). This flora spread across North America to Europe via a land
bridge between the continents.
About eleven million years ago, during the Miocene epoch, a marked
change in vegetation occurred, with the appearance of grasses and their
244
subsequent spread to grassy plains and prairies. The appearance of this

widespread flora supported the development and evolution of herbivo-
rous mammals.
The Quaternary period (1.8 mya to present) began with continental gla-
ciation in northwest Europe, Siberia, and North America. This glaciation
affected land vegetation, with plants migrating north and south as a re-
sponse to glacial and interglacial fluctuations. Pollen grains and spores
document the presence of Aceraceae (maple), hazel, and Fraxinus (ash)
during interglacial periods.
Final migrations of plant species at the close of the last ice age (about
eleven thousand years ago), formed the modern geographical distribution
of land plants. Some areas, such as mountain slopes or islands, have un-
usual distribution of plant species as a result of their isolation from the
global plant migrations.
Mariana Louise Rhoades
See also: Adaptations and their mechanisms; Adaptive radiation; Biomes:

determinants; Biomes: types; Coevolution; Colonization of the land; Con-
vergence and divergence; Dendrochronology; Evolution: definition and
theories; Evolution: history; Extinctions and evolutionary explosions; Iso-
lating mechanisms; Natural selection; Nonrandom mating, genetic drift,
and mutation; Paleoecology; Punctuated equilibrium vs. gradualism; Spe-
ciation.

Cleal, Christopher J., and Barry A. Thomas. Plant Fossils: The History of Land
Vegetation. Suffolk, England: Boydell Press, 1999.
Stewart, Wilson N., and Gar A. Rothwell. Paleobotany and the Evolution of
Plants. New York: Cambridge University Press, 1993.
245
EXTINCTIONS AND
EVOLUTIONARY EXPLOSIONS
Types of ecology: Evolutionary ecology; Paleoecology; Population

ecology; Speciation
The history of life has been punctuated by episodes of great change, some marked by
the loss of large numbers of organisms, others by explosive development. Explana-
tions proposed for these fluctuations have a bearing on current extinction levels
and the extent to which they can be controlled.
E xtinction of species is a continuous process, and evidence of its occur-

rence abounds in the fossil record. It has been estimated that marine
species persist for about four million years, which translates into an overall
loss of about two or three species each year. This is the “background” ex-
tinction rate, and it is balanced by speciation events that result in the devel-
opment of new species. Mass extinctions are events during which the rate
of extinction rises dramatically above this background rate, and a number
of these have been recognized in the Phanerozoic era. In each of these
events, at least 40 percent of the genera of shallow marine organisms were
eliminated. Using statistical methods, it has been estimated that at least 65
percent of species became extinct at each of these events, with 77 percent
being eliminated at the event at the end of the Cretaceous period and 95
percent at the event at the end of the Permian period. These mass extinc-
tions were balanced by periods of explosive development that often fol-
lowed, as organisms moved into vacant adaptive zones during periods of
adaptive radiation. The most important of these was at the base of the
Cambrian period, 544 million years ago, when all the major groups in exis-
tence originated, but other radiations occurred in the Early Triassic period
and at the start of the Tertiary period.
Causes of Mass Extinctions

Attempts to explain the causes of mass extinctions have centered on terres-
trial phenomena such as sea-level changes, climatic changes, or volcanic
activity. The sea level has shown regular fluctuations on a global level dur-
ing the Phanerozoic era, and these appear to be related to the melting or
formation of polar ice caps or to major tectonic events such as continental
splits or the collision and uplift or subsidence of ocean ridges. Extinction
events appear to be correlated mostly with periods of marine regression.
246
Extinctions and evolutionary explosions
During such a regression, the withdrawal of the ocean leaves a much

smaller habitat for shallow marine organisms. This leads to increased
crowding and competition and ultimately to an increased extinction rate.
Reduction of large terrestrial vertebrates during these regressions, as hap-
pened during the events at the end of the Permian and Cretaceous periods,
may be related to increased seasonality caused by the loss of the ameliorat-
ing influence of the shallow epicontinental seas.
It has also been shown that some extinctions are related to transgressive
events (the spread of the sea over land areas), possibly resulting from the
spread of anoxic (oxygen-poor) waters across epicontinental areas. Clima-
tic changes seem to be correlated with eustatic events (worldwide changes
in sea level), and the evidence implicating temperature as the main cause
of extinctions seems weak. For example, the most important extinction
event at the end of the Permian period occurred at a time of climatic ame-
lioration marked by the disappearance of the Gondwanaland ice sheet.
Volcanic activity has been presented as a possible cause of the extinctions
that occurred at the end of the Cretaceous period. The Deccan Traps of
northern India were erupting at that time and would have produced large
quantities of volatile emissions that could have resulted in global cooling,
ozone-layer depletion, and changes in ocean chemistry. However, no evi-
dence exists as yet for the involvement of volcanic activity in other extinc-
tion events.
Although various extraterrestrial causes for mass extinction events
have been suggested in the past, these ideas have gained greater credence
since the publication in the early 1980’s of work by Luis and Walter
Alvarez, who ascribe the end-Cretaceous extinction event to the effects of
the impact of a large bolide, or extraterrestrial object, perhaps ten kilome-
ters in diameter. The impact of such a large object would have resulted in
some months of darkness because of the global dust clouds generated, and
this would have halted photosynthesis and resulted in the collapse of both
terrestrial and marine food chains. Although cold would initially have ac-
companied the darkness, greenhouse effects and global warming would
follow as atmospheric gases and water vapor trapped infrared energy ra-
diating from the earth. Physical evidence for an impact rests on the pres-
ence in the period boundary layers of high concentrations of iridium and
other elements generally rare at the earth’s surface but abundant in aster-
oids. In addition, these layers often contain shocked quartz grains, other-
wise found only in impact craters and at nuclear test sites, and microtectites,
glassy droplets formed by impact. Although the evidence for extraterres-
trial impacts having caused the other major extinction events is slight, this
causal factor has been linked with the apparently regular 26-million-year
247
periodicity exhibited by extinctions. Scientists suggest that the regular pas-

sage of an unidentified planetary body by the Oort Cloud of comets and
the subsequent perturbation could result in increased asteroid impacts and
extinction events on earth.
Historical Mass Extinctions

The first mass extinction event that can be recognized in the fossil record
occurred in the Middle Vendian period, about 650 million years ago, when
microorganisms underwent a severe decline. This event has been linked to
climatic cooling related to glaciation. The extinction in the Late Ordovician
period was a major event in which 22 percent of marine families became
extinct. As there were two main pulses of extinction and no iridium anom-
aly was found, an extraterrestrial cause seems unlikely. However, sea-level
and temperature changes have been cited as likely causes. In addition, bio-
logically toxic bottom waters might have been brought to the surface dur-
ing periods of climatic change. The event at the end of the Devonian period
had a devastating effect on brachiopods, which lost about 86 percent of
genera, and on reef-building organisms such as corals. Shallow-water fau-
nas were most severely affected; only 4 percent of species survived, al-
though 40 percent of deeper-water species did, and cool-water faunas also
survived better. This event has been linked to a significant drop in global
temperatures of unknown cause.
The mass extinction event at the end of the Permian period was the
most severe of the Phanerozoic era and resulted in the extinction of up
to 95 percent of all marine invertebrate species. On land, amphibians and
mammal-like reptiles were both badly affected, and plant diversity fell by
50 percent. No iridium anomaly was found, and the most likely expla-
nation is climatic instability caused by continental amalgamation and
the simultaneous occurrence of marine regressions. These occurrences
would have disrupted food webs on a major scale. The event that occurred
at the end of the Triassic period was much less severe but still involved
extensive reductions in marine invertebrates and reptiles. On land, a ma-
jor faunal turnover took place. Primitive amphibians, early reptile groups,
and mammal-like reptiles died out and were replaced by advanced rep-
tiles and mammals. No evidence of an impact event has been found, and
the extinctions are generally correlated with widespread marine regres-
sions.
The extinction that took place at the end of the Cretaceous period has
become the most hotly debated, in large part because of the bolide impact
hypothesis. Although the broad pattern of extinction among marine or-
ganisms is known, the detailed picture only encompasses microorganisms
248
such as planktonic foraminifera and calcareous nannoplankton. Study of

the ranges of these microorganisms shows that the extinctions occurred
over an extended period, starting well before and finishing well after the
boundary. Although much has been made of the extinction of ammonites
at the end of the Cretaceous period, there are too few ammonite-bearing
sections to show if it was gradual or abrupt. On land, evidence of an in-
crease in the population of ferns just above the boundary suggests the pres-
ence of wildfires, as ferns are usually the first plants to recolonize an area
devastated by fire. However, in many sections, a return of the Cretaceous
vegetation is seen above the fern increase, indicating little extinction.
Post-Extinction Recoveries
Among the vertebrates, a picture of gradual change is seen for mammals,
with drastic reductions occurring only in the marsupials. The boundary
also does not seem to have been a barrier for turtles, crocodiles, lizards,
and snakes, all of which came through virtually unscathed. The dinosaurs
did become extinct, and much argument has centered on whether this was
abrupt or occurred after a slow decline. In this context, it must be noted
that there is only one area where a dinosaur-bearing sedimentary transi-
tion across the boundary can be seen, and that is in Alberta, Canada, and
the northwestern United States. Records of dinosaurs in this area during
the upper part of the Cretaceous period show a gradual decline in diver-
sity, with a drop from thirty to seven genera over the last eight million
years. Although explanations of the extinction of dinosaurs have ranged
from mammals eating their eggs to terminal allergies caused by the rise of
flowering plants to the current ideas about bolide impacts, the answer may
be climate related. A major regression of the oceans occurred at this point,
resulting in a drop in mean annual temperatures and an increase in season-
ality. The bolide impact may have served as the death blow to taxa (ani-
mals in classification groups) that were already declining.
The main period of evolutionary expansion in the Phanerozoic era is at
the base of the Cambrian period, 544 million years ago. Termed the “Cam-
brian explosion,” it marks the development of all the modern phyla of or-
ganisms, and as many as one hundred phyla may have existed during the
Cambrian period. This period seems to have lasted only about 5 million
years, and the subsequent history of animal life consists mainly of varia-
tions on the anatomical themes developed during this short period of in-
tense creativity. This period is represented in the fossil record by the re-
markably well-preserved Burgess Shale fauna of British Columbia, which
has been extensively described, and faunas of similar age from China and
Greenland. Why the Cambrian explosion could establish all major anatom-
249
ical designs so quickly is not clear. Some scientists believe that the lack of
complex organisms before the explosion had left large areas of ecological
space open, and when experimentation took place, particularly with the
advent of hard skeletons, any novelty could find a niche. Also, the earliest
multicellular organisms may have maintained a genetic flexibility that be-
came greatly reduced as organisms became locked into stable and success-
ful designs. Why some of the innovations were successful in the long term
and others were not is unknown, as no recognized traits unite the success-
ful taxa. It has even been suggested that success may be due to no more
than the luck of the draw.
In contrast, the recoveries after the major extinctions at the end of the
Permian and Cretaceous periods did not result in the development of new
phyla. The earliest Triassic ecosystems were more vacant than at any time
since the Cambrian period, yet no new phyla or classes appear in the Trias-
sic period. This suggests that despite the overwhelming nature of the ex-
tinctions, the pattern was insufficient to permit major morphological inno-
vations, in part probably because no adaptive zone was entirely vacant.
Hence, despite the fact that the mass extinction at the end of the Permian
period triggered an explosion in marine diversity described as the Meso-
zoic marine revolution, persisting species may have limited the success of
broad evolutionary jumps.
Reading the Fossil Records

All understanding of extinction events or of evolutionary explosions de-
pends on the fossil record. The study of the diversity of organisms through
time—the number of different types of organisms that occur at a particular
time and place—is therefore very important. The basic data consists of
compilations of extinctions of taxa plotted against similar compilations of
origination of taxa. Periods when either extinction or origination were un-
usually high show as peaks or troughs on a graph. Unfortunately, biases in
the preservation, collection, and study of fossils have conspired to obscure
patterns of change in diversity.
The geological history of patterns of diversity are obscured by a vari-
ety of filters, many of which are sampling biases that cause the observed
fossil record to differ from the actual history of the biosphere. The most
severe bias is the loss of sedimentary rock volume and area as the age of the
record increases because the volume and area correlate strongly with the
diversity of organisms described from a stratigraphic interval. The qual-
ity of the record also tends to fall with increasing age because the rocks
are exposed to changes that may destroy the fossils they contain. The dif-
ferences in levels of representation among the paleoenvironments in the
250
stratigraphic record also influences the composition of the fossil record; for
example, shallow-marine faunas are much better represented than are ter-
restrial faunas.
Diversity patterns are studied at a variety of levels, from the species up-
ward, that vary in their quality and inclusiveness. A basic problem is that
many of the processes that are of interest occur at the species level or even
below it, but the biases of the fossil record mean that data is best at higher
levels. Diversity of shallow-marine organisms for the Phanerozoic era can-
not be read directly at the species level because the record is too fragmen-
tary. The record at the family level is much more complete because the
preservation of one species in a family allows the family to be recorded. For
this reason, paleodiversity studies are often conducted at the family level.
However, higher taxon diversity is a poor predictor of species diversity.
For example, an analysis of the mass extinction at the end of the Permian
period indicates that the 17 percent reduction in marine orders and 52 per-
cent reduction in marine families probably represents a 95 percent reduc-
tion in the number of species. Another problem with the study of fossils is
that soft-bodied and poorly skeletonized groups may leave little or no re-
cord. It has generally been assumed that the ratio of heavily skeletonized
to non-skeletonized species has remained approximately constant through
the Phanerozoic era; however, there is no data to support this and some ev-
idence that skeletons have become more robust through time in response
to newly evolving predators. The net result of these biases is severe. Only
10 percent of the skeletonized marine species of the geologic past and far
fewer of the soft-bodied species are known.
Despite these problems, it has been possible to show that diversity of or-
ganisms has varied in a number of ways during the Phanerozoic era. Tabu-
lations of classes, orders, and families have been used to show that there
were significant periods of increased extinction or increased evolutionary
rates. One of the most important uses of this data has been the tabulation at
the family level that appears to show a regular periodicity of about 26 mil-
lion years for extinction events and that has been used to support ideas
about periodic extraterrestrial events. However, although fluctuations oc-
curred, it has also been possible to show that the number of marine orders
increased rapidly to the Late Ordovician period and has remained approx-
imately constant since then.
The Ebb and Flow of Life on Earth

Mass extinctions and evolutionary explosions are the opposite faces of
the pattern of diversity of organisms through time. During periods of mass
extinction, the diversity of organisms on earth has dropped drastically
251
and in some cases entire lineages have been wiped out. Evolutionary ex-
plosions, on the other hand, resulted in enormous innovation, particu-
larly at the beginning of the Cambrian period, and the development of
new variations on established morphotypes (animal and plant forms and
structures) later in the geologic record. Understanding the processes that
caused these events is of major importance because people have reached
the point where they are capable of influencing their environment in dras-
tic ways.
Studies of extinction events have shown that they have a variety of
causes, some of which appear to be environmental changes brought about
by natural processes while others may be the result of extraterrestrial
forces. The most severe of these extinction events occurred at the end of the
Permian period, 245 million years ago, and resulted in the loss of up to 95
percent of marine invertebrate species. The cause of this extinction appears
primarily to be that continents were amalgamating and oceans were re-
treating, which resulted in a major reduction in the habitat of shallow-
marine organisms. Terrestrial habitats were also affected as the increase in
continental area and loss of the ameliorating effect of extensive areas of
shallow ocean brought about climatic changes. Although climatic changes
are thought to be the main culprit in the majority of extinction events, some
scientists believe that large bolides, or extraterrestrial bodies, struck the
earth with such force as to create major changes in the environment that
significantly reduced diversity. This theory has enjoyed the most popular-
ity as the explanation for the event at the end of the Cretaceous period, dur-
ing which the dinosaurs became extinct, but evidence for an extraterres-
trial body’s involvement in any of the other events is slight.
Whatever the cause, environmental change that results in habitat reduc-
tion is the main reason for species decline. As humans have risen to domi-
nance over other species, the extinction rate has accelerated, and in the last
half-century, this rate has climbed considerably above natural attrition as
populations have increased and habitats have been altered. Although the
levels of extinction have not yet reached those recorded during major ex-
tinction events of the past, some scientists believe people may be facing an
ecological disaster. A better understanding of the processes surrounding
past extinction events and the rebounds that followed them will help peo-
ple prepare for and deal with the future.
David K. Elliott

chronology; Development and ecological strategies; Evolution: definition
252
and theories; Evolution: history; Evolution of plants and climates; Gene

flow; Genetic drift; Genetically modified foods; Isolating mechanisms;
Natural selection; Nonrandom mating, genetic drift, and mutation; Paleo-
ecology; Population genetics; Punctuated equilibrium vs. gradualism;
Speciation; Species loss.

Allen, Keith C., and Derek E. Briggs, eds. Evolution and the Fossil Record.
Washington, D.C.: Smithsonian Press, 1989.
Alvarez, W. T. Rex and the Crater of Doom. Princeton, N.J.: Princeton Univer-
sity Press, 1997.
Archibald, J. D. Dinosaur Extinction and the End of an Era. New York: Colum-
Bakker, Robert T. The Dinosaur Heresies. New York: William Morrow, 1986.
Briggs, Derek E., and Peter R. Crowther, eds. Palaeobiology: A Synthesis. Ox-
ford, England: Blackwell Scientific Publications, 1990.
Donovan, Stephen K., ed. Mass Extinctions: Processes and Evidence. London:
Belhaven Press, 1989.
Drury, Stephen. Stepping Stones: Evolving the Earth and Its Life. New York:
Oxford University Press, 1999.
Erwin, Douglas H. The Great Paleozoic Crisis. New York: Columbia Univer-
sity Press, 1993.
Frankel, Charles. The End of the Dinosaurs: Chicxulub Crater and Mass Extinc-
tions. New York: Cambridge University Press, 1999.
Gould, Stephen J. Wonderful Life: The Burgess Shale and the Nature of History.
New York: W. W. Norton, 1989.
Leakey, Richard E., and Roger Lewin. The Sixth Extinction: Patterns of Life
and the Future of Humankind. New York: Doubleday, 1995.
McGhee, George R. The Late Devonian Mass Extinction. New York: Colum-
McMenamin, Mark A., and Dianna L. McMenamin. The Emergence of Ani-
mals: The Cambrian Breakthrough. New York: Columbia University Press,
1989.
Martin, Paul S., and Richard G. Klein, eds. Quaternary Extinctions: A Prehis-
toric Revolution. Tucson: University of Arizona Press, 1984.
Muller, Richard. Nemesis: The Death Star. New York: Weidenfeld &
Nicolson, 1988.
Officer, Charles, and Jake Page. The Great Dinosaur Extinction Controversy.
Boston: Addison-Wesley, 1996.
Raup, David M. Extinction: Bad Genes or Bad Luck? New York: W. W.
Norton, 1991.
253
Runnegar, Bruce, and James W. Schopf, eds. Major Events in the History of
Life. Boston: Jones and Bartlett, 1992.
Stanley, Steven M. Extinction. New York: Scientific American Library, 1987.
Stearns, Beverly Petersen, and Stephen C. Stearns. Watching, from the Edge
of Extinction. New Haven, Conn.: Yale University Press, 1999.
Ward, Peter D., and Don Brownlee. Rare Earth: Why Complex Life Is Uncom-
mon in the Universe. New York: Copernicus, 2000.
254
FOOD CHAINS AND WEBS
Types of ecology: Community ecology; Ecoenergetics
The concept of food chains and webs allows ecologists to interconnect the organ-
isms living in an ecosystem and to trace mathematically the flow of energy from
plants through animals to decomposers.
T he food chain concept provides the basic framework for production bi-
ology and has major implications for agriculture, wildlife biology, and
calculating the maximum amount of life that can be supported on the
earth. As early as 1789, naturalists such as Gilbert White described the
many sequences of animals eating plants and themselves being eaten by
other animals. However, the use of the term “food chain” dates from 1927,
when Charles Elton described the implications of the food chain and food
web concept in a clear manner. His solid exposition advanced the study of
two important biological concepts: the complex organization and interre-
latedness of nature, and energy flow through ecosystems.
Food Chains in Ecosystem Description

Stephen Alfred Forbes, founder of the Illinois Natural History Survey, con-
tended in 1887 that a lake comprises a system in which no organism or pro-
cess can be understood unless its relationship to all the parts is understood.
Forty years later, Elton’s food chains provided an accurate way to diagram
these relationships. Because most organisms feed on several food items,
food chains were cross-linked into complex webs with predictive power.
For instance, algae in a lake might support an insect that in turn was food
for bluegill. If unfavorable conditions eliminated this algae, the insect
might also disappear. However, the bluegill, which fed on a wider range of
insects, would survive because the loss of this algae merely increases the
pressure on the other food sources. This detailed linkage of food chains ad-
vanced agriculture and wildlife management and gave scientists a solid
overview of living systems. When Arthur George Tansley penned the term
ecosystem in the 1930’s, it was food-chain relationships that described
much of the equilibrium of the ecosystem.
Today most people still think of food chains as the basis for the “balance
of nature.” This phrase dates from the controversial 1960 work of Nelson
G. Hairston, Frederick E. Smith, and Lawrence B. Slobodkin. They pro-
posed that if only grazers and plants are present, grazing limits the plants.
With predators present, however, grazers are limited by predation, and the
255
Food chains and webs
plants are free to grow to the limits of the nutrients available. Such expla-
nations of the “balance of nature” were commonly taught in biology books
throughout the 1960’s and 1970’s.
Food Chains in Production Biology

Elton’s explanation of food chains came just one year after Nelson Transeau
of Ohio State University presented his calculations on the efficiency with
which corn plants converted sunlight into plant tissue. Ecologists traced
this flow of stored chemical energy up the food chain to herbivores that ate
plants and on to carnivores that ate herbivores. Food chains therefore un-
dergirded the new “production biology” that placed all organisms at vari-
ous trophic levels and calculated the extent to which energy was lost or
preserved as it passed up the food chain.
With data accumulating from many ecologists, Elton extended food
chains into a pyramid of numbers. The food pyramid, in which much plant
tissue supports some herbivores that are in turn eaten by fewer carnivores,
is still referred to as an Eltonian pyramid. In 1939 August Thienemann
added decomposers to reduce unconsumed tissues and return the nutri-
ents of all levels back to the plants. Early pyramids were based on the
amount of living tissues, or biomass.
Calculations based on the amount of chemical energy at each level, as
measured by the heat released when food is burned (calories), provided
even more accurate budgets. Because so much energy is lost at each stage
in a food chain, it became obvious that this inefficiency was the reason food
chains are rarely more than five or six links long and why large, fierce ani-
mals are uncommon. It also became evident that because the earth inter-
cepts a limited amount of sunlight energy per year, there is a limit on the
amount of plant life—and ultimately upon the amount of animal life and
decomposers—that can be fed. Food chains are also important in the ac-
counting of carbon, nitrogen, and water cycling.
Value of Food Chains in Environmental Science

Unlike calories, which are dramatically reduced at each step in a food
chain, some toxic substances become more concentrated as the molecules
are passed along. The concentration of molecules along the food chain was
first noticed by the Atomic Energy Commission, which found that radioac-
tive iodine and strontium released in the Columbia River were concen-
trated in tissue of birds and fish. However, the pesticide DDT provided the
most notorious example of biological magnification: DDT was found to be
deposited in animal body fat in ever-increasing concentrations as it moved
up the food chain to ospreys, pelicans, and peregrine falcons. High levels
256
Food chains and webs
of DDT in these birds broke down steroid hormones and interfered with
eggshell formation.
Because humans are omnivores, able to feed at several levels on the
food chain (that is, both plants and other animals), it has been suggested
that a higher world population could be supported by humans moving
down the food chain and becoming vegetarians. A problem with this argu-
ment is that much grazing land worldwide is unfit for cultivation, and
therefore the complete cessation of pig or cattle farming does not necessar-
ily free up substantial land to grow crops.
While the food chain and food web concepts are convenient theoretical
ways to summarize feeding interactions among organisms, real field situa-
tions have proved far more complex and difficult to measure. Animals of-
ten switch diet between larval and adult stages, and they are often able to
shift food sources widely. It is often difficult to draw the boundaries of
food chains and food webs.
John Richard Schrock
See also: Balance of nature; Biomass related to energy; Geochemical cy-

cles; Herbivores; Hydrologic cycle; Nutrient cycles; Omnivores; Phyto-
plankton; Rain forests and the atmosphere; Trophic levels and ecological
niches.

Colinvaux, Paul A. Why Big Fierce Animals Are Rare. New York: Penguin
Books, 1990.
Elton, Charles. Animal Ecology. New York: October House, 1966.
Golley, Frank B. A History of the Ecosystem Concept in Ecology. New Haven,
Quamman, David. The Song of the Dodo: Island Biogeography in an Age of Ex-
tinction. New York: Simon & Schuster, 1997.
“Something’s Fishy.” Science News 146, no. 1 (July 2, 1994): 8.
Svitil, Kathy A. “Collapse of a Food Chain.” Discover 16, no. 7 (July, 1995):
36.
257
FOREST FIRES
Whether natural or caused by humans, fires destroy life and property in forest-
lands but are also vital to the health of forests.
E vidence of forest fires is routinely found in soil samples and tree bor-
ings. The first major North American fires in the historical record were
the Miramichi and Piscataquis fires of 1825. Together, they burned 3 mil-
lion acres in Maine and New Brunswick. Other U.S. fires of significance
were the Peshtigo fire in 1871, which raged over 1.28 million acres and took
fourteen hundred human lives in Wisconsin; the fire that devastated north-
ern Idaho and northwestern Montana in 1910 and killed at least seventy-
nine firefighters; and a series of fires that joined forces to sweep across one-
third of Yellowstone National Park in 1988.
Fire Behavior
Fires need heat, fuel, and oxygen. They spread horizontally by igniting
particles at their edge. At first, flames burn at one point, then move out-
ward, accumulating enough heat to keep burning on their own. Topogra-
phy and weather affect fire behavior. Fires go uphill faster than downhill
because warm air rises and preheats the uphill fuels. Vegetation on south-
and west-facing slopes receives maximal sunlight and so is drier and burns
more easily. Heat is pulled up steep, narrow canyons, as it is up a chimney,
increasing heat intensity. For several reasons, only one-third of the vegeta-
tion within a large fire usually burns. This mosaic effect may be caused by
varied tree species that burn differently, old burns that stop fire, strong
winds that blow the fire to the leeward side of trees, and varied fuel mois-
ture.
Ecological Benefits
Some plant species require very high temperatures for their seed casings to
split for germination. After fire periodically sweeps through the forest,
seeds will germinate. Other species, such as the fire-resistant ponderosa
pine, require a shallow layer of decaying vegetable matter in which to root.
Fires burn excess debris and small trees of competing species and leave an
open environment suitable for germination. Dead material on the forest
floor is processed into nutrients more quickly by fire than by decay, and in
258
Forest fires
a layer of rich soil, plants will sprout within days to replace those de-
stroyed in the fire.
Ecological Destruction
Erosion is one of the devastating effects of a fire. If the fuels burn hot, tree
oils and resins can be baked into the soil, creating a hard shell that will not
absorb water. When it rains, the water runoff gathers mud and debris, cre-
ating flash floods and extreme stream sedimentation. Culverts and storm
drains fill with silt, and streams flood and change course. Fish habitat is de-
stroyed, vegetation sheltering stream banks is ripped away, and property
many miles downstream from the forest is affected.
When a fire passes through timber it generally leaves pockets of green,
although weakened, stands. Forest pests, such as the bark beetle, are at-
tracted to the burned trees and soon move to the surviving trees, weaken-
ing them further. Healthy trees outside the burn area may also fall to pest
infestation unless the burned trees are salvaged before pests can take hold.
During the twentieth century,

low fires that otherwise would
have burned through the forest
at ground level every five to
twenty-five years were
suppressed. As a result, the
natural cycle of frequent fires
moving through an area was
broken, and when fire did erupt,
accumulated kindling burned
hot and fast, exploding into
devastating crown fires and
completely destroying the local
community’s habitat.
(PhotoDisc)
259
Forest fires
The ash and smoke from hot, fast-burning forest fires can be transported
for miles, affecting air quality many miles from the actual fire.
Ecological Impact of Fire Suppression

One of the early criteria of forest management was fire protection. In the
second quarter of the twentieth century, lookout towers, firebreaks, and
trails were built to locate fires as quickly as possible. Low fires that other-
wise would have burned through the forest at ground level and cleared out
brush every five to twenty-five years were suppressed. As a result, the nat-
ural cycle of frequent fires moving through an area was broken. Fallen
trees, needles, cones, and other debris collected as kindling on the forest
floor, rather than being incinerated every few years.
It took foresters and ecologists fifty years to realize that too much fire
suppression was as bad as too little. Infrequent fires cause accumulated
kindling to burn hot and fast and explode into treetops. The result is a dev-
astating crown fire, a large fire that advances as a single front. Burning em-
bers of seed cones and sparks borne by hot, strong winds created within
the fire are tossed into unburned areas to start more fires.
Prescribed Burns
In the 1970’s prescribed burning was added to forest management tech-
niques used to keep forests healthy. Fires set by lightning are allowed to
burn when the weather is cool, the area isolated, and the risk of the fire ex-
ploding into a major fire low. More than 70 percent of prescribed burning
takes place in the southeastern states, where natural fires burn through an
area more frequently than in the West.
In some areas, prescribed fires are set in an attempt to re-create the natu-
ral sequence of fire. In Florida, prescribed burns provide wildlife habitat
by opening up groves to encourage healthy growth. Other fires start acci-
dentally but are allowed to burn until they reach a predetermined size.
Although a prescribed fire is an attempt to duplicate natural fire, it is
not as efficient, because private and commercial property within the fire
path must be protected. Once a fire has occurred, burned timber deterio-
rates quickly, either through insect infestation or blueing—a mold that
stains the wood. Private landowners can move quickly to salvage fire-
damaged trees and plant new seedlings to harness erosion. On federal
land, regulations governing the salvage of trees can delay logging of the
burned snags until deterioration makes it uneconomical to harvest them.
Causes
Forest fires may be caused by natural events or human activity. Most natu-
260
Forest fires
ral fires are started by lightning strikes. Dozens of strikes can be recorded
from one lightning storm. When a strike seems likely, fire spotters watch
for columns of smoke, and small spotter planes will fly over the area, look-
ing for smoke. Many of the small fires simply smoulder and go out, but if
the forest is dry, multiple fires can erupt from a single lightning storm.
The majority of forest fires are human-caused, and most are the result of
carelessness rather than arson. Careless campers may leave a campsite
without squelching their campfire completely, and winds may then whip
the glowing embers into flames. A smoker may toss a cigarette butt from a
car window. Sparks from a flat tire riding on the rim may set fire to vegeta-
tion alongside the highway. The sun shining through a piece of broken
glass left by litterers may ignite dry leaves.
Fire Fighting
In fire fighting, bulldozers are used to cut fire lines ahead of the approach-
ing fire, and fuels between fire lines and the fire are backburned. Helicop-
ters and tanker planes drop water with a fire-retardant additive, or benton-
ite, a clay, at the head of the fire to smother fuels. Firefighters are equipped
with fire shelters in the form of aluminized pup tents, which they can pull
over themselves if a fire outruns them. Despite technological advances,
one of the best tools for fighting fires—along with the shovel—remains the
pulaski, a combination ax and hoe, first produced commercially in 1920.
This tool, in the hands of on-the-ground firefighters, is used to cut fire
breaks and to throw dirt on smoldering debris.
Public Policy and Public Awareness

Since the early twentieth century, forest fires have engendered public pol-
icy in the United States. In the aftermath of major fires in 1903 and 1908 in
Maine and New York, state fire organizations and private timber protec-
tive associations were formed to provide fire protection. These, in turn,
contributed to the Weeks Act of 1911, which permitted cooperative fire
protection between federal and state governments.
People who make their homes in woodland settings in or near forests
face the danger of forest fire, and government agencies provide informa-
tion to help people safeguard themselves and their property. Homes near
forests should be designed and landscaped with fire safety in mind, using
fire-resistant or noncombustible materials on the roof and exterior. Land-
scaping should include a clear safety zone around the house. Hardwood
trees, less flammable than conifers, and other fire-resistant vegetation
should be planted.
J. A. Cooper
261
Forest fires
See also: Communities: structure; Erosion and erosion control; Forest

management; Forests; Mediterranean scrub; Mountain ecosystems; Old-
growth forests; Paleoecology; Rain forests; Savannas and deciduous tropi-
cal forests; Taiga; Trophic levels and ecological niches.

Fuller, Margaret. Forest Fires: An Introduction to Wildland Fire Behavior, Man-
agement, Firefighting, and Prevention. New York: Wiley, 1991.
Johnson, Edward A. Fire and Vegetation Dynamics: Studies from the North
American Boreal Forest. New York: Cambridge University Press, 1992.
Kasischke, Eric S., and Brian J. Stocks, eds. Fire, Climate Change, and Carbon
Cycling in the Boreal Forest. New York: Springer, 2000.
Pringle, Laurence P. Fire in the Forest: A Cycle of Growth and Renewal. New
York: Simon and Schuster, 1995.
Pyne, Stephen J. Fire in America: A Cultural History of Wildland and Rural
Fire. 2d ed. Seattle: University of Washington Press, 1997.
Whelan, Robert J. The Ecology of Fire. New York: Cambridge University
Press, 1995.
262
FOREST MANAGEMENT
Forest management includes reforestation programs as well as techniques to man-

age logging practices, provide grazing lands, support mining operations, main-
tain infrastructure networks, or slow the destruction of rain forests.
F orests provide lumber for buildings, wood fuel for cooking and heat-
ing, and raw materials for making paper, latex rubber, resin, dyes, and
essential oils. Forests are also home to millions of plants and animal species
and are vital in regulating climate, purifying the air, and controlling water
runoff. A 1993 global assessment by the United Nations Food and Agricul-
ture Organization (FAO) found that three-fourths of the forests in the
world still have some tree cover, but less than one-half of these have intact
forest ecosystems. Deforestation is occurring at an alarming rate, and man-
agement practices are being sought to try to halt this destruction.
Thousands of years ago, forests and woodlands covered almost 15 bil-
lion acres of the earth. Approximately 16 percent of the forests have been
cleared and converted to pasture, agricultural land, cities, and nonpro-
ductive land. The remaining 11.4 billion acres of forests cover about 30 per-
cent of the earth’s land surface. Clearing forests has severe environmental
consequences. It reduces the overall productivity of the land, and nutrients
and biomass stored in trees and leaf litter are lost. Soil once covered with
plants, leaves, and snags becomes prone to erosion and drying. When for-
ests are cleared, habitats are destroyed and biodiversity is greatly dimin-
ished. Destruction of forests causes water to drain off the land instead of
being released into the atmosphere by transpiration or percolation into
groundwater. This can cause major changes in the hydrologic cycle and ul-
timately in the earth’s climate. Because forests remove a large amount of
carbon dioxide from the air; the clearing of forests causes more carbon di-
oxide to remain, thus upsetting the delicate balance of atmospheric gases.
Rain Forests
Rain forests provide habitats for at least 50 percent (some estimates are as
high as 90 percent) of the total stock of plant, insect, and other animal spe-
cies on earth. They supply one-half of the world’s annual harvest of hard-
wood and hundreds of food products, such as chocolate, spices, nuts, coffee,
and tropical fruits. Tropical rain forests also provide the main ingredients
in 25 percent of prescription and nonprescription drugs, as well as 75 per-
263
Forest management
cent of the three thousand plants identified as containing chemicals that

fight cancer. Industrial materials, such as natural latex rubber, resins, dyes,
and essential oils, are also harvested from tropical forests.
Tropical forests in Asia, Africa, and Latin America are rapidly being
cleared to produce pastureland for large cattle ranches, establish logging
operations, construct large plantations, grow narcotic plants, develop min-
ing operations, or build dams to provide power for mining and smelting
operations. In 1985 the FAO’s Committee on Forest Development in the
Tropics developed the Tropical Forestry Action Plan to combat these prac-
tices, develop sustainable forest methods, and protect precious ecosys-
tems. Fifty nations in Asia, Africa, and Latin America have adopted the
plan.
Management Techniques
Several management techniques have been successfully applied to slow
the destruction of tropical forests. Sustainable logging practices and refor-
estation programs have been established on lands that allow timber cut-
ting, with complete bans of logging on virgin lands. Certain regions have
set up extractive reserves to protect land for the native people who live in
the forest and gather latex rubber and nuts from mature trees. Sections of
some tropical forests have been preserved as national reserves, which at-
tract tourists while preserving trees and biodiversity.
Developing countries have been encouraged to protect their tropical
forests by using a combination of debt-for-nature swaps and conservation
easements. In debt-for-nature swaps, tropical countries act as custodians
of the tropical forest in exchange for foreign aid or relief from debt. Conser-
vation easement involves having another country, private organization, or
consortium of countries compensate a tropical country for protecting a
specific habitat.
Another management technique involves putting large areas of the for-
est under the control of indigenous people who use slash-and-burn agri-
culture (also known as swidden or milpa agriculture). This traditional,
productive form of agriculture follows a multiple-year cycle. Each year
farmers clear a forest plot of several acres in size to allow the sun to pene-
trate to the ground. Leaf litter, branches, and fallen trunks are burned, leav-
ing a rich layer of ashes. Fast-growing crops, such as bananas and papayas,
are planted and provide shade for root crops, which are planted to anchor
the soil. Finally, crops such as corn and rice are planted. Crops mature in a
staggered sequence, thus providing a continuous supply of food. Use of
mixed perennial polyculture helps prevent insect infestations, which can
destroy monoculture crops. After one or two years, the forest begins to
264
Forest management
take over the agricultural plot. The farmers continue to pick the perennial
crops but essentially allow the forest to reclaim the plot for the next ten to
fifteen years before clearing and planting the area again. Slash-and-burn
agriculture can, however, post hazards: A drought in Southeast Asia in
1997 caused such fires to burn for months when monsoon rains did not ma-
terialize, polluting the air and threatening the health of millions of Indone-
sians. In 1998, previous abuse of the technique resulted in flooding and
mudslides in Honduras after the onset of Hurricane Mitch.
U.S. Forest Management

Forests cover approximately one-third of the land area of the continental
United States and comprise 10 percent of the forests in the world. Only
about 22 percent of the commercial forest area in the United States lies
within national forests. The rest is primarily managed by private compa-
nies that grow trees for commercial logging. Land managed by the U.S.
Forest Service provides inexpensive grazing lands for more than three mil-
lion cattle and sheep every year, supports multimillion-dollar mining op-
erations, and consists of a network of roads eight times longer than the U.S.
interstate highway system. Almost 50 percent of national forest land is
open for commercial logging. Nearly 14 percent of the timber harvested in
the United States each year comes from national forest lands. Total wood
production in the United States has caused the loss of more than 95 percent
of the old-growth forests in the lower forty-eight states. This loss includes
not only high-quality wood but also a rich diversity of species not found in
early-growth forests.
National forests in the United States are required by law to be managed
in accordance with principles of sustainable yield. Congress has mandated
that forests be managed for a combination of uses, including grazing, log-
ging, mining, recreation, and protection of watersheds and wildlife.
Healthy forests also require protection from pathogens and insects. Sus-
tainable forestry, which emphasizes biological diversity, provides the best
management. Other management techniques include removing only in-
fected trees and vegetation, cutting infected areas and removing debris,
treating trees with antibiotics, developing disease-resistant species of
trees, using insecticides and fungicides, and developing integrated pest
management plans.
Two basic systems are used to manage trees: even-aged and uneven-
aged. Even-aged management involves maintaining trees in a given stand
that are about the same age and size. Trees are harvested, then seeds are re-
planted to provide for a new even-aged stand. This method, which tends
toward the cultivation of a single species or monoculture of trees, empha-
265
Forest management
Environmental Effects of Select Silvicultural Methods

Long-term effects Silvicultural methods Short-term effects
Converting mixed
Soil acidification forest stands to
monoculture Increased leaching of
Fertilizing nutrients, especially
nitrogen
Forest machines
leave the residues in
heaps, often on wet
sites
Soil erosion Clear-cutting Increased water runoff
Soil scarification
Decreased number
of plant and animal Draining Increased amount of
species of the forests, organic material and
mires, and fens Short rotation intervals metals (Fe, Al, Hg) in
water and ecosystems;
Decreased number of secondary effects on fish
Decreased number of old and dead trees
forest plant and animal
species Foreign species
Source: Adapted from I. Stjernquist, “Modern Wood Fuels,” in Bioenergy and the
Environment, edited by Pasztor and Kristoferson, 1990.
sizes the mass production of fast-growing, low-quality wood (such as

pine) to give a faster economic return on investment. Even-aged manage-
ment requires close supervision and the application of both fertilizer and
pesticides to protect the monoculture species from disease and insects.
Uneven-aged management maintains trees at many ages and sizes to
permit a natural regeneration process. This method helps sustain biologi-
cal diversity, provides for long-term production of high-quality timber,
allows for an adequate economic return, and promotes a multiple-use ap-
proach to forest management. Uneven-aged management also relies on se-
lective cutting of mature trees and reserves clear-cutting for small patches
of tree species that respond favorably to such logging methods.
Harvesting Methods
The use of a particular tree-harvesting method depends on the tree species
involved, the site, and whether even-aged or uneven-aged management is
being applied. Selective cutting is used on intermediate-aged or mature
266
Forest management
trees in uneven-aged forests. Carefully selected trees are cut in a prescribed

stand to provide for a continuous and attractive forest cover that preserves
the forest ecosystem.
Shelterwood cutting involves removing all the mature trees in an area
over a period of ten years. The first harvest removes dying, defective, or
diseased trees. This allows more sunlight to reach the healthiest trees in the
forest, which will then cast seeds and shelter new seedlings. When the
seedlings have turned into young trees, a second cutting removes many of
the mature trees. Enough mature trees are left to provide protection for the
younger trees. When the young trees become well established, a third cut-
ting harvests the remaining mature trees, leaving an even-aged stand of
young trees from the best seed trees to mature. When done correctly, this
method leaves a natural-looking forest and helps reduce soil erosion and
preserve wildlife habitat.
Seed-tree cutting harvests almost every tree at one site, with the excep-
tion of a few high-quality seed-producing and wind-resistant trees, which
will function as a seed source to generate new crops. This method allows a
variety of species to grow at one time and aids in erosion control and wild-
life conservation.
Clear-cutting removes all the trees in a single cutting. The clear-cut may
involve a strip, an entire stand, or patches of trees. The area is then re-
planted with seeds to grow even-aged or tree-farm varieties. More than
two-thirds of the timber produced in the United States, and almost one-
third of the timber in national forests, is harvested by clear-cutting. A clear-
cut reduces biological diversity by destroying habitat. It can make trees in
bordering areas more vulnerable to winds and may take decades to regen-
erate.
Forest Fires
Forest fires can be divided into three types: surface, crown, and ground
fires. Surface fires tend to burn only the undergrowth and leaf litter on the
forest floor. Most mature trees easily survive, as does wildlife. These fires
occur every five years or so in forests with an abundance of ground litter
and help prevent more destructive crown and ground fires. Such fires can
release and recycle valuable mineral nutrients, stimulate certain plant
seeds, and help eliminate insects and pathogens.
Crown fires are very hot fires that burn both ground cover and tree tops.
They normally occur in forests that have not experienced fires for several
decades. Strong winds allow these fires to spread from deadwood and
ground litter to treetops. They are capable of killing all vegetation and
wildlife, leaving the land prone to erosion.
267
Forest management
Ground fires are more common in northern bogs. They can begin as sur-
face fires but burn peat or partially decayed leaves below the ground sur-
face. They can smolder for days or weeks before anyone notices them, and
they are difficult to douse.
Natural forest fires can be beneficial to some plant species, including the
giant sequoia and the jack pine trees, which release seeds for germination
only after being exposed to intense heat. Grassland and coniferous forest
ecosystems that depend on fires to regenerate are called fire climax ecosys-
tems. They are managed for optimum productivity with prescribed fires.
The Society of American Foresters has begun advocating a concept
called new forestry, in which ecological health and biodiversity, rather
than timber production, are the main objectives of forestry. Advocates of
new forestry propose that any given site should be logged only every 350
years, wider buffer zones should be left beside streams to reduce erosion
and protect habitat, and logs and snags should be left in forests to help re-
plenish soil fertility. Proponents also wish to involve private landowners
in the cooperative management of lands.
Toby R. Stewart and Dion Stewart
See also: Communities: structure; Conservation biology; Deforestation;

Erosion and erosion control; Forest fires; Forests; Grazing and overgrazing;
Integrated pest management; Multiple-use approach; Old-growth forests;
Reforestation; Restoration ecology; Species loss; Sustainable development;
Urban and suburban wildlife; Wildlife management.

Davis, Kenneth P. Forest Management. New York: McGraw-Hill, 1996.
Davis, Lawrence S., and K. Norman Johnson. Forest Management. 3d ed.
New York: McGraw-Hill, 1987.
Hunter, Malcolm L., Jr. Wildlife, Forests, and Forestry. Englewood Cliffs, N.J.:
McNeely, Jeffrey A. Conserving the World’s Biological Diversity. Washington,
D.C.: WRI, 1990.
Nyland, Ralph D. Silviculture: Concepts and Applications. 2d ed. Boston:
McGraw-Hill, 2002.
Robbins, William G. American Forestry. Lincoln: University of Nebraska
Press, 1985.
Smith, David M. The Practice of Silviculture. 9th ed. New York: Wiley, 1997.
Spurr, Stephen H., and Burton V. Barnes. Forest Ecology. 4th ed. New York:
Wiley, 1998.
268
FORESTS
Forests are complex ecosystems in which trees are the dominant type of plant.
There are three main forest biomes: tropical, temperate, and boreal.
B oth humans and animals depend on forests for food, shelter, and other
resources. Forests once covered much of the world and are still found
from the equator to the Arctic regions. A forest may vary in size from only a
few acres to thousands of square miles, but generally any natural area in
which trees are the dominant type of plant can be considered a forest. For a
plant to be called a tree, the standard definition requires that the plant
must attain a mature height of at least 8 feet (about 3 meters), have a woody
stem, and possess a distinct crown. Thus, size makes roses shrubs and ap-
ples trees, even though apples and roses are otherwise close botanical rela-
tives. Foresters generally divide the forests of the world into three general
categories: tropical, temperate, and boreal.
Tropical Rain Forest

The tropical rain forest is a forest consisting of a dizzying variety of trees,
shrubs, and other plants that remain green year-round. The growth is lush
and usually includes both a dense canopy formed by the crowns of the
largest trees and a thick understory of smaller trees and shrubs. Growth is
often continuous, rather than broken into periods of dormancy and active
growth, so that fruiting trees are occasionally seen bearing blossoms and
mature fruit simultaneously.
Temperate Forest
The temperate forest lies between the tropical forest and the boreal, or
northern, forest. The forests of the Mediterranean region of Europe as well
as the forests of the southern United States are temperate forests. Trees in
temperate forests can be either deciduous or coniferous. Although conifer-
ous trees are generally thought of as evergreen, the distinction between
types is actually based on seed production and leaf shape. Coniferous
trees, such as spruces, pines, and hemlocks, produce seeds in cones and
have needle-like leaves. Deciduous trees, such as maples, poplars, and
oaks, have broad leaves and bear seeds in other ways. Some conifers, such
as tamarack, do change color and drop their needles in the autumn, while
269
Forests
some deciduous trees, particularly in the southerly regions of the temper-

ate forest, are evergreen.
Deciduous trees are also referred to as hardwoods, while conifers are
softwoods, a classification that refers more to the typical density of the
wood than to how difficult it is to nail into it. Softwoods are lower in den-
sity and will generally float in water while still green. Hardwoods are
higher in density on average and will sink.
Like tropical forests, temperate forests can be quite lush. While the
dominant species vary from area to area, depending on factors such as soil
types and available rainfall, a dense understory of shade-tolerant species
often thrives beneath the canopy. Thus, a mature temperate forest may
have thick stands of rhododendrons 20 to 30 feet (6 to 9 meters) high thriv-
ing in the shade of 80-foot (24-meter) oaks and tulip poplar. As the temper-
ate forest approaches the edges of its range and the forest makes the transi-
tion to boreal, the understory thins out, disappearing almost completely or
consisting only of low shrubs. Even in temperate forests, the dominant
species may prevent an understory from forming. Stands of southern
loblolly pine, for example, often have a parklike feel, as the thick mulch
created by fallen needles chokes out growth of other species.
Boreal Forest
The boreal forest, which lies in a band across the northern United States,
Canada, northern Europe, and northern Asia, is primarily a coniferous for-
est. The dominant species are trees such as white spruce, hemlock, and
white pine. Mixed stands of northern hardwoods, such as birch, sugar ma-
ple, and red oak, may be found along the southern reaches of the boreal
forest. As the forest approaches the Arctic, trees are fewer in type, becom-
ing primarily spruce, birch, and willows, and smaller in size. The under-
story is generally thin or nonexistent, consisting of seedlings of shade-
tolerant species, such as maple, and low shrubs. Patches of boreal-type
forest can be found quite far south in higher elevations in the United States,
such as the mountains of West Virginia. The edge of the temperate forest
has crept steadily northward following the retreat of the glaciers at the end
of the Ice Age twenty thousand years ago.
Forest Ecology and Resources

In all three types of forest a complex system of interrelationships governs
the ecological well-being of the forest and its inhabitants. Trees and ani-
mals have evolved to fit into particular environmental niches. Some wild-
life may need one resource provided by one species of tree in the forest
during one season and a resource provided by another during a different
270
Forests
Forest Areas by Region

temperate/boreal
North America, 13.2%
(457 million hectares; Latin America and
1,129 million acres) the Caribbean, 27.5%
(950 million hectares;
Europe, 4.2% 2,347 million acres)
361 million acres)
former USSR, 23.6% Africa, 15.1%

(816 million hectares; (520 million hectares;
2,016 million acres) 1,284 million acres)
Asia/Oceania, 16.4%
1,396 million acres)
1995: total area = 3,454 million hectares; 8,533 million acres
Source: Data are from United Nations Food and Agriculture Organization (FAOSTAT
Database, 2000)
time of year, while other animals become totally dependent on one specific
tree. Whitetail deer, for example, browse on maple leaves in the summer,
build reserves of fat by eating acorns in the fall, and survive the winter by
eating evergreens. Deer are highly adaptable in contrast with other spe-
cies, such as the Australian koala, which depends entirely on eucalyptus
leaves for its nutritional needs. Just as the animals depend on the forest, the
forest depends on the animals to disperse seeds and thin new growth. Cer-
tain plant seeds, in fact, will not sprout until being abraded as they pass
through the digestive tracts of birds.
Humans also rely on the forest for food, fuel, shelter, and other prod-
ucts. Forests provide wood for fuel and construction, fibers for paper,
and chemicals for thousands of products often not immediately recog-
nized as deriving from the forest, such as plastics and textiles. In addi-
tion, through the process of transpiration, forests regulate the climate by
releasing water vapor into the atmosphere while removing harmful car-
bon compounds. Forests play an important role in the hydrology of a
watershed. Rain that falls on a forest will be slowed in its passage down-
hill and is often absorbed into the soil rather than running off into rivers
271
Forests
and lakes. Thus, forests can moderate the effects of severe storms, reducing
the dangers of flooding and preventing soil erosion along stream and river
banks.
Threats to the Forest

The primary threat to the health of forests around the world comes from
humans. As human populations grow, three types of pressure are placed
on forests. First, forests are cleared to provide land for agriculture or for the
construction of new homes. This process has occurred almost continuously
in the temperate regions for thousands of years, but it did not become com-
mon in tropical regions until the twentieth century. Often settlers level the
forest and burn the fallen trees to clear land for farming (slash-and-burn
agriculture) without the wood itself being utilized in any way. Tragically,
the land thus exposed can become infertile for farming within a few years.
After a few years of steadily diminishing crops, the land is abandoned.
With the protective forest cover removed, it may quickly become a barren,
eroded wasteland.
Second, rising or marginalized populations in developing nations often
depend on wood or charcoal as their primary fuel for cooking and for
home heat. Forests are destroyed as mature trees are removed for fuel
wood faster than natural growth can replace them. As the mature trees dis-
appear, younger and younger growth is also removed, and eventually the
forest is gone completely.
Finally, growing populations naturally demand more products derived
from wood, which can include everything from lumber for construction to
chemicals used in cancer research. Market forces can drive forest products
companies to harvest more trees than is ecologically sound as stockholders
focus on short-term individual profits rather than long-term environmen-
tal costs. The challenge to foresters, ecologists, and other scientists is to de-
vise methods that allow humanity to continue to utilize the forest re-
sources needed to survive without destroying the forests as complete and
healthy ecosystems.
See also: Biomes: types; Deforestation; Erosion and erosion control; Forest
fires; Forest management; Grazing and overgrazing; Mountain ecosys-
tems; Multiple-use approach; Old-growth forests; Paleoecology; Rain for-
ests; Rain forests and the atmosphere; Reforestation; Restoration ecology;
Savannas and deciduous tropical forests; Species loss; Sustainable de-
velopment; Taiga; Trophic levels and ecological niches; Tundra and high-
altitude biomes; Urban and suburban wildlife; Wildlife management.
272
Forests

Holland, Israel I. Forests and Forestry. 5th ed. Danville, Ill.: Interstate, 1997.
Kimmins, J. P. Balancing Act: Environmental Issues in Forestry. 2d ed. Vancou-
ver: UBC Press, 1997.
Page, Jake. Forest. Rev. ed. Alexandria, Va.: Time-Life Books, 1987.
Walker, Laurence C. The Southern Forest. Austin: University of Texas Press,
1991.
273
GENE FLOW
Types of ecology: Community ecology; Evolutionary ecology;
Population ecology; Speciation
Gene flow represents a recurrent exchange of genes between populations. This ex-
change results when immigrants from one population interbreed with members of
another.
C harles Darwin published On the Origin of Species by Means of Natural

Selection in 1859. Since then, scientists have modified and added new
concepts to the theory of evolution by natural selection. One of those con-
cepts, which was only dimly understood in Darwin’s lifetime, is the impor-
tance of genetics in evolution, especially the concepts of migration and
gene flow.
Genes
Genes are elements within the cells of a living organism that control the
transmission of hereditary characteristics by specifying the structure of a
particular protein or by controlling the function of other genetic material.
Within any species, the exchange of genes via reproduction is constant
among its members, ensuring genetic similarity. If a new gene or combina-
tion of genes appears in the population, it is rapidly dispersed among all
members of the population through inbreeding. New alleles (forms of a
gene) may be introduced into the gene pool of a breeding population (thus
contributing to the evolution of that species) in two ways: mutation and
migration. Gene flow is integral to both processes.
A mutation occurs when the DNA code of a gene becomes modified so
that the product of the gene will also be changed. Mutations occur con-
stantly in every generation of every species. Most of them, however, are ei-
ther minor or detrimental to the survival of the individual and thus are of
little consequence. A very few mutations may prove valuable to the sur-
vival of a species and are spread to all of its members by migration and
gene flow.
Separation and Migration

In nature, gene flow occurs on a more or less regular basis between demes,
geographically isolated populations, races, and even closely related spe-
cies. Gene flow is more common among the adjacent demes of one species.
The amount of migration between such demes is high, thus ensuring that
274
Gene flow
their gene pools will be similar. This sort of gene flow contributes little to
the evolutionary process, as it does little to alter gene frequencies or to con-
tribute to variation within the species.
Much more significant for the evolutionary process is gene flow be-
tween two populations of a species that have not interbred for a prolonged
period of time. Populations of a species separated by geographical barriers
(as a result, for example, of seed dispersal to a distant locale) often develop
very dissimilar gene combinations through the process of natural selec-
tion. In isolated populations, dissimilar alleles become fixed or are present
in much different frequencies. When circumstances do permit gene flow to
occur between populations, it results in the breakdown of gene complexes
and the alteration of allele frequencies, thereby reducing genetic differ-
ences in both. The degree of this homogenization process depends on the
continuation of interbreeding among members of the two populations
over extended periods of time.
Hybridization
The migration of a few individuals from one breeding population to an-
other may, in some instances, also be a significant source of genetic varia-
tion in the host population. Such migration becomes more important in the
evolutionary process in direct proportion to the differences in gene fre-
quencies—for example, the differences between distinct species. Biologists
call interbreeding between members of separate species hybridization.
Hybridization usually does not lead to gene exchange or gene flow, be-
cause hybrids are not often well adapted for survival and because most are
sterile. Nevertheless, hybrids are occasionally able to breed (and produce
fertile offspring) with members of one or sometimes both the parent spe-
cies, resulting in the exchange of a few genes or blocks of genes between
two distinct species. Biologists refer to this process as introgressive hybrid-
ization. Usually, few genes are exchanged between species in this process,
and it might be more properly referred to as “gene trickle” rather than gene
flow.
Introgressive hybridization may, however, add new genes and new
gene combinations, or even whole chromosomes, to the genetic architec-
ture of some species. It may thus play a role in the evolutionary process, es-
pecially in plants. Introgression requires the production of hybrids, a rare
occurrence among highly differentiated animal species but quite common
among closely related plant species. Areas where hybridization takes place
are known as contact zones or hybrid zones. These zones exist where pop-
ulations overlap. In some cases of hybridization, the line between what
constitutes different species and what constitutes different populations of
275
Gene flow
the same species becomes difficult to draw. The significance of introgres-

sion and hybrid zones in the evolutionary process remains an area of some
contention among life scientists.
Speciation
Biologists often explain, at least in part, the poorly understood phenome-
non of speciation through migration and gene flow—or rather, by a lack
thereof. If some members of a species become geographically isolated from
the rest of the species, migration and gene flow cease. Such geographic iso-
lation can occur, for example, when populations are separated by water (as
occurs on different islands or other landmasses) or valleys (different hill-
sides). The isolated population will not share in any mutations, favorable
or unfavorable, nor will any mutations that occur among its own members
be transmitted to the general population of the species. Over long periods
of time, this genetic isolation will result in the isolated population becom-
ing so genetically different from the parent species that its members can no
longer produce fertile progeny should one of them breed with a member of
the parent population. The isolated members will have become a new spe-
cies, and the differences between them and the parent species will continue
to grow as time passes. Scientists, beginning with Darwin, have demon-
strated that this sort of speciation has occurred on the various islands of the
world’s oceans and seas.
Paul Madden
See also: Adaptive radiation; Clines, hybrid zones, and introgression;

Convergence and divergence; Evolution: definition and theories; Evolu-
tion of plants and climates; Extinctions and evolutionary explosions; Ge-
netic diversity; Genetic drift; Isolating mechanisms; Natural selection;
Nonrandom mating, genetic drift, and mutation; Punctuated equilibrium
vs. gradualism; Speciation.

Christiansen, Freddy B. Population Genetics of Multiple Loci. New York:
Wiley, 2000.
Endler, John A. Geographic Variation, Speciation, and Clines. Princeton, N.J.:
Princeton University Press, 1977.
Foster, Susan A., and John A. Endler, eds. Geographic Variation in Behavior:
Perspectives on Evolutionary Mechanisms. New York: Oxford University
Press, 1999.
Leapman, Michael. The Ingenious Mr. Fairchild: The Forgotten Father of the
Flower Garden. New York: St. Martin’s Press, 2001.
276
Gene flow
Mousseau, Timothy A., Barry Sinervo, and John A. Endler, eds. Adaptive
Genetic Variation in the Wild. New York: Oxford University Press, 2000.
Real, Leslie A., ed. Ecological Genetics. Princeton, N.J.: Princeton University
Press, 1994.
Stuessy, Tod F., and Mikio Ono, eds. Evolution and Speciation of Island Plants.
New York: Cambridge University Press, 1998.
277
GENETIC DIVERSITY
Types of ecology: Community ecology; Ecosystem ecology; Population
ecology; Restoration and conservation ecology
Genetic diversity includes the inherited traits encoded in the DNA of all living or-
ganisms and can be examined on four levels: among species, among populations
(in communities), within populations, and within individuals. Populations with
higher levels of diversity are better able to adapt to changes in the environment and
are more resistant to the deleterious effects of inbreeding.
G enetic diversity is the most fundamental level of biological diversity

because genetic material is responsible for the variety of life. For new
species to form, genetic material must change. Changes in the inherited
properties of populations occur deterministically through gene flow (mat-
ing between individual organisms representing formerly separated popu-
lations) and through natural or artificial selection (which occurs when
some types of individuals breed more successfully than others). Change
can also occur randomly through mutations or genetic drift (when the rela-
tive proportions of genes change by chance in small populations). Popula-
tions with higher levels of diversity tend to do better—to have more sur-
vival options—as surroundings change than do populations (particularly
smaller ones) with lower levels of genetic diversity.
Preservation Efforts
Conservation efforts directed at maintaining genetic diversity involve
both germ plasm preservation (germ plasm kept in a steady state for peri-
ods of time) and germ plasm conservation (germ plasm kept in a natural,
evolving state). The former usually involves ex situ laboratory techniques
in which genetic resources are removed from their natural habitats. They
include seminatural strategies such as botanical gardens, arboreta, nurser-
ies, zoos, farms, aquaria, and captive fisheries, as well as completely artifi-
cial methods such as seed reserves or “banks,” microbial cultures (preserv-
ing bacteria, fungi, viruses, and other microorganisms), tissue cultures of
parts of plants and animals (including sperm storage), and gene libraries
(involving storage and replication of partial segments of plant or animal
DNA, or deoxyribonucleic acid).
Conservation areas are the preferred in situ (at the natural or original
place) means of protecting genetic resources. Ideally these include preserv-
ing the number and relative proportions of species and the genetic diver-
278
Genetic diversity
sity they represent, the physical features of the habitat, and all ecosystem
processes. It is not always enough, however, to maintain the ecosystem
which the threatened species inhabits. It is sometimes necessary to take an
active interventionist position in order to save a species. Controversial
strategies can include reintroduction of captive species into the wild,
sometimes after they have been genetically manipulated. Direct manage-
ment of the ecosystem may also be attempted by either lessening human
exploitation and interference or by reducing the number of natural preda-
tors or competitors. However, management of a specific conservation area
varies in terms of what is valued and how preservation is accomplished.
Crop Diversity
One area of keen interest that illustrates the issues involved with the pres-
ervation of any kind of genetic diversity is how to preserve crop germ
plasm. Largely conserved in gene banks, crop germ plasm was historically
protected by farmers who selected for success in differing environments
and other useful traits. Traditionally cultivated varieties (landraces) diver-
sified as people spread into new areas. Colonial expansion produced new
varieties as farmers adapted to new conditions and previously separated
plant species interbred; other species were lost when some societies de-
clined and disappeared.
By the early 1900’s field botanists and agronomists were expressing
concern about the rapidly escalating loss of traditionally cultivated variet-
ies. This loss accelerated after the 1940’s as high yielding hybrids of cereal
and vegetable crops replaced local landraces. Wild relatives of these land-
races are also disappearing as their habitats are destroyed through human
activity. Gene banks preserve both kinds of plants because, as argued by
Nikolai I. Vavilov in 1926, crop plant improvement can best be accom-
plished by taking advantage of these preserved genetic stocks. Vavilov
also noted that genetic variation for most cultivated species was concen-
trated in specific regions, his “centers of diversity,” most of which are re-
gions where crop species originated.
The vulnerability to parasites and climate of an agriculture that relies on
one or a few varieties of crops necessitates the maintenance of adequate re-
serves of genetic material for breeding. In addition to the preservation of
species known to be useful, many people advocate preservation of wild
species for aesthetic reasons as well as for their unknown future potential.
Maintenance of Productivity
Farmers in developed nations change crop varieties every four to ten years
in order to maintain consistent levels of food production. This necessitates
279
Genetic diversity
an ongoing search for new breeds with higher yields and an ability to with-
stand several environmental challenges, including resistance to multiple
pests and drought. Over time, older varieties mutate, become less popular
at the marketplace, or are unable to adapt to new conditions. However,
farmers from poorer nations are not always able to take advantage of the
new breeds or afford the expensive support systems, including chemical
fertilizers. Moreover, not all types of crops have benefited equally from
conservation efforts.
Another tension between the world’s poor and rich nations concerns
ownership of genetic diversity. The Convention on Biodiversity, signed by
167 nations in 1992, states that genetic materials are under the sovereign
control of the countries in which they are found. This policy is particularly
controversial regarding medicinal plants, because “biodiversity prospect-
ing” for new drugs has economically benefited either individuals or corpo-
rations based in the developed countries.
Joan C. Stevenson
See also: Adaptive radiation; Clines, hybrid zones, and introgression;

Convergence and divergence; Evolution: definition and theories; Evolu-
tion of plants and climates; Extinctions and evolutionary explosions; Gene
flow; Isolating mechanisms; Natural selection; Punctuated equilibrium vs.
gradualism; Speciation.

Hawkes, John G. The Diversity of Crop Plants. Cambridge, Mass.: Harvard
Fundamentals of Conservation Biology. Cambridge, Mass.: Blackwell Science,
1996.
Krattiger, Anatole F., et al., eds. Widening Perspectives on Biodiversity. Geneva,
Switzerland: International Academy of the Environment, 1994.
Orians, Gordon H., et al., eds. The Preservation and Valuation of Biological Re-
sources. Seattle: University of Washington Press, 1990.
280
GENETIC DRIFT
Types of ecology: Evolutionary ecology; Population ecology
Genetic drift refers to random changes in the genetic composition of a population.

It is one of the evolutionary forces that cause biological evolution, the others being
natural selection, mutation, and migration, or gene flow.
D rift occurs because the genetic variants, or alleles, present in a popula-

tion are a random sample of the alleles that adults in the previous
generation would have been predicted to pass on, where predictions are
based on expected migration rates, expected mutation rates, and the direct
effects of alleles on fitness. If this sample is small, then the genetic composi-
tion of the offspring population may deviate substantially from expecta-
tion, just by chance. This deviation is called genetic drift. Drift becomes in-
creasingly important as population size decreases. The key feature of drift
that distinguishes it from the other evolutionary forces is the unpredictable
direction of evolutionary change.
Anything that generates fitness variation among individuals (that is,
variation in the ability of individuals to survive and reproduce) will in-
crease the magnitude of drift for all genes that do not themselves cause the
fitness variation. Because of their indeterminate growth, plants often vary
greatly in reproductive potential because of local environmental variation,
and this magnifies genetic drift. For example, the magnitude of drift in
most annual plants is more than doubled by size variation among adults.
This makes sense if one considers that larger individuals contribute a
larger number of offspring to the next generation, so any alleles they carry
will tend to be overrepresented.
Fitness variation caused by selection will also increase the magnitude of
drift at any gene not directly acted upon by the selection. If an individual
has high fitness because it possesses one or more favorable alleles, then all
other alleles it possesses will benefit. This is called genetic hitchhiking.
This is a potent source of evolution because the direction of change at a
hitchhiking gene will remain the same for multiple generations. However,
it is not possible to predict in advance what that direction will be because
where and when a favorable mutation will occur cannot be predicted.
The opportunities for drift to occur are greatly influenced by gene flow.
Most terrestrial plants are characterized by highly localized dispersal.
Thus, even in large, continuous populations, the pool of potential mates
for an individual, and the pool of seeds that compete for establishment at a
281
Genetic drift
site, are all drawn from a small number of nearby individuals known as the
neighborhood. If the neighborhood is sufficiently small, genetic drift will
have a significant impact on its genetic composition.
For these and other reasons, population size alone is not sufficient to
predict the magnitude of drift. The effective size of a population, Ne , is a
number that is directly related to the magnitude of drift through a simple
equation. Thus, Ne incorporates all characteristics of a population that in-
fluence drift.
Loss of Variability
The long-term consequence of drift is a loss of genetic variation. As alleles
increase and decrease in frequency at random, some will be lost. In the ab-
sence of mutation and migration, such losses are permanent. Eventually,
only one allele remains at each gene, which is said to be fixed. Thus, all else
being equal, smaller populations are expected to harbor less genetic varia-
tion than larger populations.
An important way in which different plant populations are not equal is
in their reproductive systems. With self-fertilization (selfing), or asexual
reproduction, genetic hitchhiking becomes very important. In the extreme
cases of 100 percent selfing or 100 percent asexual reproduction, hitchhik-
ing will determine the fates of most alleles. Thus, as a new mutation
spreads or is eliminated by selection, so too will most or all of the other al-
leles carried by the individual in which the mutation first arose. This is
called a selective sweep, and the result is a significant reduction in genetic
variation. Which alleles will be swept to fixation or elimination cannot be
predicted in advance, so the loss of variation reflects a small Ne. Consistent
with this expectation, most populations of flowering plants that reproduce
partly or entirely by selfing contain significantly less genetic variation than
populations of related species that do not self-fertilize.
Extinction
Mutations that decrease fitness greatly outnumber mutations that increase
fitness. In a large population in which drift is weak, selection prevents
most such mutations from becoming common. In very small populations,
however, alleles that decrease fitness can drift to fixation, causing a de-
crease in average fitness. This is one manifestation of a phenomenon called
inbreeding depression. In populations with very small Ne , this inbreeding
depression can be significant enough to threaten the population with ex-
tinction. If a population remains small for many generations, mean fitness
will continue to decline as new mutations become fixed by drift. When fit-
ness declines to the point where offspring are no longer overproduced,
282
Genetic drift
population size will decrease further. Drift then becomes stronger, muta-
tions are fixed faster, and the population heads down an accelerating tra-
jectory toward extinction. This is called mutational meltdown.
Creative Potential
By itself, drift cannot lead to adaptation. However, drift can enhance the
ability of selection to do so. Because of diploidy and sexual recombination,
some types of mutations, either singly or in combinations, will increase fit-
ness when common but not when rare. Genetic drift can cause such genetic
variants to become sufficiently common for selection to promote their fixa-
tion. A likely example is the fixation of new structural arrangements of
chromosomes that occurred frequently during the diversification of flow-
ering plants. New chromosome arrangements are usually selected against
when they are rare because they disrupt meiosis and reduce fertility. The
initial spread of such a mutation can therefore only be caused by strong ge-
netic drift, either in an isolated population of small effective size or in a
larger population divided into small neighborhoods.
John S. Heywood
See also: Biodiversity; Clines, hybrid zones, and introgression; Extinctions

and evolutionary explosions; Gene flow; Genetic diversity; Isolating mecha-
Population genetics; Punctuated equilibrium vs. gradualism; Speciation.

Denny, Mark, and Steven Gaines. Chance in Biology: Using Probability to Ex-
plore Nature. Princeton, N.J.: Princeton University Press, 2000.
Futuyma, Douglas J. Evolutionary Biology. 3d ed. Sunderland, Mass.: Sinauer
Associates, 1998.
283
GENETICALLY MODIFIED FOODS
Types of ecology: Agricultural ecology; Chemical ecology;
Ecotoxicology; Evolutionary ecology
Applications of genetic engineering in agriculture and the food industry could in-
crease world food supplies, reduce environmental problems associated with food
production, and enhance the nutritional values of certain foods. However, these
benefits are countered by food-safety concerns, the potential for ecosystem disrup-
tion, and fears of unforeseen consequences resulting from altering natural selec-
tion.
H umans rely on plants and animals as food sources and have long
used microbes to produce foods such as cheese, bread, and fer-
mented beverages. Conventional techniques such as cross-hybridization,
production of mutants, and selective breeding have resulted in new varie-
ties of crop plants or improved livestock with altered genetics. However,
these methods are relatively slow and labor-intensive, are generally lim-
ited to intraspecies crosses, and involve a great deal of trial and error.
Transgenic Technology
Recombinant DNA techniques, which manipulate cells’ deoxyribonucleic
acid (DNA), were developed in the 1970’s and enabled researchers to make
specific, predetermined genetic changes in a variety of organisms. Because
the technology also allows for the transfer of genes across species and
kingdom barriers, an infinite number of novel genetic combinations are
possible. The first animals and plants containing genetic material from
other organisms (transgenics) were developed in the early 1980’s. By 1985
the first field trials of plants engineered to be pest-resistant were con-
ducted. In 1990 the U.S. Food and Drug Administration (FDA) approved
chymosin as the first substance produced by modified organisms to be
used in the food industry for dairy products such as cheese. That same year
the first transgenic cow was developed to produce human milk proteins
for infant formula. The well-publicized Flavr Savr tomato, modified to de-
lay ripening and rotting, obtained FDA approval in 1994.
Goals and Uses

By the mid-1990’s, more than one thousand genetically modified crop
plants were approved for field trials. The goals for altering food crop
plants by genetic engineering fall into three main categories: to create
284
Genetically modified foods
plants that can adapt to specific environmental conditions to make better

use of agricultural land, increase yields, or reduce losses; to increase nutri-
tional value or flavor; and to alter harvesting, transport, storage, or pro-
cessing properties for the food industry. Many genetically modified crops
are sources of ingredients for processed foods and animal feed.
Herbicide-resistant plants, such as the Roundup Ready soybean, can be
grown in the presence of glyphosphate, an herbicide that normally de-
stroys all plants with which it comes in contact. Beans from these plants
were approved for food-industry use in several countries, but there has
been widespread protest by activists such as Jeremy Rifkin and envi-
ronmental organizations such as Greenpeace. Frost-resistant fruit contain-
ing a fish antifreeze gene, insect-resistant plants with a bacterial gene that
encodes for a pesticidal protein (Bacillus thuringiensis), and a viral disease-
Genetically Modified Crop Plants Unregulated by

the U.S. Department of Agriculture
Crop Patent Holder Genetically Engineered Trait
Canola AgrEvo herbicide tolerance
Corn AgrEvo herbicide tolerance
Ciba-Geigy insect resistance
DeKalb herbicide tolerance; insect resistance
Monsanto herbicide tolerance; insect resistance
Northrup King insect resistance
Cotton Calgene herbicide tolerance; insect resistance
DuPont herbicide tolerance
Monsanto herbicide tolerance; insect resistance
Papaya Cornell virus resistance
Potato Monsanto insect resistance
Squash Asgrow virus resistance
Upjohn virus resistance
Soybean AgrEvo herbicide tolerance
DuPont altered oil profile
Monsanto herbicide tolerance
Tomato Agritope altered fruit ripening
Calgene altered fruit ripening
Monsanto altered fruit ripening
Zeneca altered chemical content in fruit
Source: U.S. Department of Agriculture and Plant Health Inspection Service (APHIS).
285
resistant squash are examples of other genetically modified food crops that
have undergone field trials.
Scientists have also created plants that produce healthier unsaturated
fats and oils rather than saturated ones. Genetic engineering has yielded
coffee plants whose beans are caffeine-free without processing and toma-
toes with altered pulp content for improved canned products. Genetically
modified microbes are used for the production of food additives such as
amino acid supplements, sweeteners, flavors, vitamins, and thickening
agents. In some cases, these substances had to be obtained from slaugh-
tered animals. Altered organisms are also used for improving fermenta-
tion processes in the food industry.
Ecological Implications
Food safety and quality are at the center of the genetically modified food
controversy. Concerns include the possible introduction of new toxins or
allergens into the diet and changes in the nutrient composition of foods.
Proponents argue that food sources could be designed to have enhanced
nutritional value.
A large percentage of crops worldwide are lost each year to drought,
temperature extremes, and pests. Plants have already been engineered to
exhibit frost, insect, disease, and drought resistance. Such alterations would
increase yields and allow food to be grown in areas that are currently too
dry or infertile, positively impacting the world food supply.
Environmental problems such as deforestation, erosion, pollution, and
loss of biodiversity have all resulted, in part, from conventional agricul-
tural practices. Use of genetically modified crops could allow better use of
existing farmland and lead to a decreased reliance on pesticides and fertil-
izers. However, critics fear the creation of unpredictable ecological con-
sequences as well. For example, “superweeds”—either the engineered
plants or new plant varieties formed by the transfer of recombinant genes
conferring various types of resistance to wild species—might compete
with valuable plants and have the potential to destroy ecosystems and
farmland unless stronger poisons were used for eradication. The transfer
of genetic material to wild relatives (outcrossing, or “genetic pollution”)
might also lead to the development of new plant diseases.
Diane White Husic
See also: Biopesticides; Erosion and erosion control; Grazing and over-
grazing; Integrated pest management; Multiple-use approach; Pesticides;
Rangeland; Slash-and-burn agriculture; Soil; Soil contamination; Species
loss.
286

American Chemical Society. Genetically Modified Foods: Safety Issues. Wash-
ington, D.C.: Author, 1995.
Anderson, Luke. Genetic Engineering, Food, and Our Environment. White
River Junction, Vt.: Chelsea Green, 1999.
Paredes-Lopez, Octavio, ed. Molecular Biotechnology for Plant Food Produc-
tion. Lancaster, Pa.: Technomic, 1999.
Rissler, Jane. The Ecological Risks of Engineered Crops. Cambridge, Mass.:
MIT Press, 1996.
Shannon, Thomas A., ed. Genetic Engineering: A Documentary History. West-
port, Conn.: Greenwood Press, 1999.
Yount, Lisa. Biotechnology and Genetic Engineering. New York: Facts on File,
2000.
287
GEOCHEMICAL CYCLES
Types of ecology: Ecoenergetics; Ecosystem ecology; Global ecology
Geochemical cycles refer to the movement, or cycling, of elements through ecosys-

tems and the biosphere. Both biotic (living) and abiotic (nonliving) components
participate in these cycles, and their interdependency means that a change in one
component of such a cycle will have an impact on all other components.
G eochemical cycles are generally considered to be those involving nu-

trient elements utilized by organisms in various ecosystems. Cycling
involves both biological and chemical processes. While nearly all natural el-
ements could be considered as being cycled through both abiotic and living
systems, certain elements are most commonly described in such systems.
These include carbon, nitrogen, phosphorus, and a variety of lesser elements
(including iron, sulfur, and trace elements such as copper and mercury).
Although the cycling of elements is often thought of as occurring in a
relatively rapid fashion, many of these elements spend long periods locked
in abiotic systems. For example, carbon may be found in materials that re-
quire millions of years to cycle through ocean sediment back into the atmo-
sphere. The fate of such elements depends on many factors, including their
chemical properties and their ability to erode or return to the atmosphere.
Some chemical elements, such as carbon, oxygen, and nitrogen, are incor-
porated into organisms from the atmosphere. Other elements, such as
phosphorus, potassium, sulfur, and iron, are found mainly in rocks and
sediments.
Carbon and Oxygen Cycles

The carbon and oxygen cycles are greatly dependent on each other. Molec-
ular oxygen, which represents approximately 20 percent of the atmo-
sphere, is used by organisms through a metabolic process called respira-
tion. In these reactions, the oxygen reacts with reduced carbon compounds
such as carbohydrates (sugars) and generates carbon dioxide (CO2).
Though carbon dioxide constitutes only a small proportion of the volume
of the atmosphere (0.04 percent), it is in this form that it is used by primary
producers such as plants. In the process of photosynthesis, utilizing sun-
light as an energy source, plants and some microorganisms bind, or fix, the
CO2, converting the carbon again into carbohydrates, resulting in growth
of the plant, or replication of the microorganism. The complex carbohy-
drates which are generated in photosynthesis serve as the food source for
288
Geochemical cycles
The Carbon Cycle

Carbon dioxide in
the atmosphere
Animal
Respiration in respiration
decomposers
Decomposition Organic
Fossil fuel Plant compounds
combustion of carbon Photosynthesis
respiration in animals
compounds in
dead organic
matter
Fossilization Feeding
Death
Organic compounds
in green plants
consumers—organisms such as animals (including humans) that eat the

plants. The carbohydrates are then broken down, regenerating carbon di-
oxide. In a sense, the combinations of respiration and photosynthesis rep-
resent the cycle of life. Approximately 70 billion metric tons of carbon diox-
ide (10 percent of the total atmospheric CO2) are fixed each year. The
concentration of carbon dioxide in the atmosphere is a factor in regulating
the temperature of Earth. Consequently, the release of large quantities of
the gas into the atmosphere through the burning of fossil fuels could po-
tentially alter the earth’s climate.
Nitrogen Cycle
Nitrogen gas (N2) represents 78 percent of the total volume of the atmo-
sphere. However, because of the extreme stability of the bond between the
two nitrogens in the gas, plants and animals are unable to use atmospheric
nitrogen directly as a nutrient. Nitrogen-fixing bacteria in the soil and in
the roots of leguminous plants (peas, clover) are able to convert the gas-
eous nitrogen into nitrites and nitrates, chemical forms that can be used by
plants. Animals then obtain nitrogen by consuming the plants. The decom-
position of nitrogen compounds results in the accumulation of ammonium
(NH4+) compounds in a process called ammonification. It is in this form
that nitrogen is commonly found under conditions in which oxygen is lim-
289
Geochemical cycles
ited. In this form, some of the nitrogen returns to the atmosphere. In the
presence of oxygen, ammonium compounds are oxidized to nitrates (nitri-
fication). Once the plant or animal has died, bacteria convert the nitrogen
back into nitrogen gas, and it returns to the atmosphere.
Phosphorus Cycle
Unlike carbon and nitrogen, which are found in the atmosphere, most of
the phosphorus required for biotic nutrition is found in mineral form.
Phosphorus is relatively water insoluble in this form; it is only gradually
dissolved in water. Available phosphorus is therefore often growth-limit-
ing in soils (it is second only to nitrogen as the scarcest of the soil nutrients).
Ocean sediments may bring the mineral to the surface through uplifting of
land, as along coastal areas, or by means of marine animals. Enzymatic
breakdown of organic phosphate by bacteria and the consumption of ma-
rine organisms by seabirds cycle the phosphorus into forms available for
use by plants. Deposition of guano (bird feces) along the American Pacific
coast has long provided a fertilizer rich in phosphorus.
Bacteria also play significant roles in the geochemical cycling of many
other elements. Iron, despite its abundance in the earth’s crust, is largely
The Phosphorus Cycle
Plant
tissues
Animal tissues
and feces
Decomposition by Urine
fungi and bacteria
Phosphates Loss in
drainage Assimilation
in solution by plant cells
Phosphates
in soil
Weathering of rock
Incorporation into sedimentary
rock; geologic uplift moves this
rock into terrestrial environments
290
Geochemical cycles
insoluble in water. Consequently, it is generally found in the form of pre-

+3
cipitates of ferric (Fe ) compounds, seen as brown deposits in water. Acids
are often formed as by-products in the formation of ferric compounds. The
bacterial oxidation of pyrite (FeS2) is a major factor in the leaching process
of iron ores and in the formation of acid mine drainage. Likewise, much of
the sulfur found in the earth’s crust is in the form of pyrite and gypsum
(CaSO4). Weathering processes return much of the sulfur to water-soluble
-2
forms; in the absence of air, the bacterial reduction of sulfate (SO4 ) to
forms such as hydrogen sulfide (H2S) allows its return to the atmosphere.
Since sulfide compounds are highly toxic to many organisms, bacterial re-
duction of sulfates is of major biogeochemical significance.
Richard Adler
See also: Balance of nature; Biomass related to energy; Communities: eco-

system interactions; Communities: structure; Food chains and webs; Her-
bivores; Hydrologic cycle; Nutrient cycles; Omnivores; Phytoplankton;
Rain forests and the atmosphere; Trophic levels and ecological niches.

Berner, Elizabeth K., and Robert A. Berner. Global Environment: Water, Air,
and Geochemical Cycles. Upper Saddle River, N.J.: Prentice Hall, 1996.
Bolin, Bert. “The Carbon Cycle.” Scientific American 223 (September, 1970).
Clark, F. E., and T. Rosswall, eds. Terrestrial Nitrogen Cycles: Processes, Eco-
system Strategies, and Management Impacts. Stockholm: Swedish Natural
Science Research Council, 1981.
Heimann, Martin. The Global Carbon Cycle. New York: Springer-Verlag,
1993.
Kasischke, Eric S., and Brian J. Stocks, eds. Fire, Climate Change, and Carbon
Cycling in the Boreal Forest. New York: Springer, 2000. Discusses the di-
rect and indirect mechanisms by which fire and climate interact to influ-
ence carbon cycling in North American boreal forests.
Lasserre, P., and J. M. Martin, eds. Biogeochemical Processes at the Land-Sea
Boundary. Amsterdam, N.Y.: Elsevier, 1986.
Pomeroy, Lawrence, ed. Cycles of Essential Elements. Stroudsburg, Pa.:
Dowden, Hutchinson, & Ross, 1974.
Tiessen, Holm, ed. Phosphorus in the Global Environment: Transfers, Cycles,
and Management. New York: Wiley, 1995.
291
GLOBAL WARMING
Type of ecology: Global ecology
Global warming is the term applied to rising global air temperatures. This rise in
temperature has the potential to cause drastic changes in climate and weather pat-
terns worldwide.
Greenhouse Effect
“Global warming” is the term for the rise in the earth’s average tempera-
ture. Scientists know that the earth’s average global temperature has been
rising since the beginning of the Industrial Revolution in the second half of
the eighteenth century. Increases in air temperature could alter precipita-
tion patterns, change growing seasons, result in coastal flooding, and turn
some areas into deserts. Scientists do not know the cause or causes of this
phenomenon but believe that it could be part of a normal climate cycle or
be caused by natural events or the activities of humankind.
When the ground is heated by sunlight, it gives off the heat as infrared
radiation. The atmosphere absorbs the infrared radiation and keeps it from
escaping into space. This is called the “greenhouse effect” because it was
once believed that the glass panes of greenhouses acted similarly to the at-
mosphere, capturing the infrared radiation given off by Earth inside the
greenhouse and not allowing it to pass through. Although it has subse-
quently been shown that greenhouses work by simply trapping the heated
air, the name has stuck.
The atmosphere eventually releases its heat into space. The amount of
heat stored in the atmosphere remains constant as long as the composition
of gases in the atmosphere does not change. Some gases, including carbon
dioxide, water vapor, and methane, store heat more efficiently than others
and are called “greenhouse gases.” If the composition of the atmosphere
changes to include more of these greenhouse gases, the air retains more
heat, and the atmosphere becomes warmer.
Global levels of greenhouse gases have been steadily increasing and in
1990 were more than 14 percent higher than they were in 1960. At the same
time, the average global temperature has also been rising. Meteorological
records show that from 1890 to the mid-1990’s, the average global tempera-
ture rose by between 0.4 and 0.7 degree Celsius. About 0.2 degree Celsius
of this temperature increase has occurred since 1950. In comparison, the
difference between the average global temperature in the 1990’s and in the
last ice age is approximately 10 degrees Celsius, and it is estimated that a
292
Global warming
drop of as little as 4 to 5 degrees Celsius could trigger the formation of con-

tinental glaciers. Therefore, the rise in average temperature is significant
and already beginning to cause some changes in the global climate. Docu-
mented changes include the melting of glaciers and rising sea levels as the
ocean gets warmer and its waters expand. Measurements of plant activity
indicate that the annual growing season has become approximately two
weeks longer in the middle latitude regions.
Effects of Global Warming

A common misunderstanding is that global warming simply means that
winters will be less cold and summers will be hotter, while everything else
U.S. Greenhouse Gas Emissions, 1990-1999

Type 1990 1994 1995 1996 1997 1998 1999
Carbon Dioxide 1,350.5 1,422.5 1,434.7 1,484.1 1,505.2 1,507.4 1,526.8
1
(carbon)
Methane Gas1 31.74 31.17 31.18 30.16 30.11 29.29 28.77
2
Nitrous Oxide 1,168 1,310 1,257 1,246 1,226 1,223 1,224
2
Chlorofluorocarbons 202 109 102 67 51 49 41
2
Halons 2.8 2.7 2.9 3.0 3.0 3.0 3.0
2
Hydrofluorocarbons
HFC-23 3.0 3.0 2.0 3.0 3.0 3.4 2.6
HFC-125 (Z) 0.3 0.5 0.7 0.9 1.1 1.3
HFC-134a 1.0 6.3 14.3 19.0 23.5 26.9 30.3
HFC-143a (Z) 0.1 0.1 0.2 0.3 0.5 0.7
Perfluorocarbons2
CF-4 3 2 2 2 2 2 2
C-2F-6 1 — 1 1 1 1 1
C-4F-10 (Z) (Z) (Z) (Z) (Z) (Z) (Z)
Sulfur hexafluoride2 1 1 2 2 2 2 1
Source: Abridged from U.S. Energy Information Administration, Emissions of Greenhouse
Gases in the United States, annual. From Statistical Abstract of the United States: 2001
(Washington, D.C.: U.S. Census Bureau, 2001).
Note: Emission estimates were mandated by Congress through Section 1605(a) of the
Energy Policy Act of 1992 (Title XVI). Gases that contain carbon can be measured either
in terms of the full molecular weight of the gas or just in terms of their carbon content.
1
In millions of metric tons.
2
In thousands of metric tons.
Z. Less than 500 metric tons.
— Represents or rounds to zero
293
Global warming
will be basically the same. Actually, the earth’s weather system is very
complicated, and higher global temperatures will result in significant
changes in weather patterns. Most changes will be observed in the middle
and upper latitudes, with equatorial regions witnessing fewer changes.
The areas that will experience most of the changes include North America,
Europe, and most of Asia. The Southern Hemisphere will experience less
severe effects because it contains more water than the Northern Hemi-
sphere does, and it takes more energy to heat water than land.
It is difficult to predict precisely what these changes will be, but obser-
vation of the changing climate and scientific studies allow researchers to
make some rough estimates of the kinds of changes Earth will experience.
Summers will be hotter, with more severe heat waves. Because hot air
holds more moisture than cool air, rain will fall less frequently in the sum-
mer. Droughts can be expected to be more common and more severe.
Through the late 1980’s and into the early 1990’s, annual temperatures
climbed higher and higher, and summer heat waves became more fre-
quent. This is a particularly troubling problem in areas where homes are
typically built without air conditioning. The air in a closed-up house dur-
ing a heat wave can reach temperatures well over 40 degrees Celsius. Be-
cause these temperatures exceed people’s normal body temperature,
which is about 36.5 degrees Celsius, it becomes very difficult for the body
to cool down. For this reason, heat waves are particularly dangerous for
the very young, the elderly, and people who are ill. More frequent heat
waves will cause increases in the use of air conditioning, which requires
more energy and will release additional greenhouse gases.
Global warming will also produce severe autumn rains. The overheated
summer air will cool in the autumn and will no longer be able to hold all of
the moisture it was storing. It will release the moisture as heavy rains,
causing flooding. This phenomenon has already been observed, but not for
a period of time that is scientifically significant. It is difficult to tell the dif-
ference between long-term changes and short-term fluctuations with only
a few years of observations.
These changing rain patterns—droughts and severe autumn storms—
will certainly have an effect on the earth’s landscape. Some areas may be
turned to deserts, and others may be transformed from plains to forests.
One of the strange aspects of global warming is that it is predicted to re-
sult in not only hotter periods during the summers but also colder periods
during the winter. It takes energy to move large cold-air masses from the
polar regions in winter, so it is possible that large winter storms will be
colder, more violent, and more frequent. This pattern has been evident
since about the mid-1970’s. Winter storms have brought record low tem-
294
Global warming
peratures and enough snow to close cities for days. However, this time
span is too short to determine whether this is a temporary phenomenon or
a trend. It is possible that some smaller, less permanent event than global
warming is responsible for the more severe winter storms. A shorter (al-
though more severe) winter may create a surge in pest populations and
diseases that are normally controlled by long winters.
Global warming will cause ocean levels to rise because water expands
when heated and also because of the melting of glaciers on Greenland and
Antarctica. The melting of the ice in the northern polar areas will not con-
tribute to rising ocean levels because that ice, unlike the ice on Greenland
and Antarctica, is already in the ocean. Just as the melting of the ice in a
drink does not cause the level of the drink to rise, the melting of the north-
ern oceanic ice sheets will not affect sea levels.
Causes of Global Warming

Although many scientists are convinced, based on the abundant evidence,
that global warming is occurring, they are less sure of why. One significant
factor, the rise in greenhouse gases, can be attributed to the activities of hu-
mankind. Burning forests to clear land and operating factories and auto-
mobiles produce carbon dioxide and water vapor. Livestock herds and rot-
ting vegetation release methane, and fertilizers used on farms also release
greenhouse gases. Power plants that consume fossil fuels such as coal, oil,
or natural gas release massive amounts of carbon dioxide into the air.
However, no one knows whether humankind’s activities are the real or
only reason for the increase in global temperature. The last ice age ended
very recently in geologic terms, and a number of changes are still taking
place as the globe recovers from the presence of huge ice sheets. It is possi-
ble that the world’s climate is still warming up from the last ice age. Vol-
canoes are another major source of greenhouse gases, and the level of vol-
canic activity has been increasing since approximately the beginning of the
Industrial Revolution.
Global warming could also be part of a natural cyclical change in the cli-
mate. Evidence indicates that the earth’s climate varies between warmer
and colder periods that last a few centuries. History provides many stories
of dramatically changing climate, including one period from 1617 to 1650
that was so unusually cold that it is called the “Little Ice Age.” Therefore,
Earth may be merely experiencing another cyclical change in its climate.
Study of Global Warming

The problem with studying global warming is that no one can be sure of its
extent, and some scientists debate whether it actually exists. They argue
295
Global warming
that the observed changes may only be a warm phase of a climatic cycle
that will make the earth warmer for a few years, then cooler for a few years.
If this is true, then the coming decades may see average temperatures lev-
eling off or even dropping.
A major source of the confusion involves the way global warming is stud-
ied. It would seem easy to record temperatures for a number of years and
then compare them. However, detailed records on the weather have not
been kept for more than a few decades in many areas. Scientists are forced to
rely on interpretations of historical accounts and the clues left in fossil rec-
ords. The analysis of tree rings, sedimentary deposits, and even very old ice
from deep within glaciers can provide data about the climate in the past.
In addition, the existing records must be reviewed carefully to identify
local changes that may not reflect global ones. For example, as towns grow
into cities, the temperature climbs simply because larger cities are warmer
than smaller ones, a phenomenon known as the urban heat island effect.
Measurements taken years ago in a more rural environment should be
lower than those taken after the population around the measuring station
increased. This problem can be overcome with balloons. By sending instru-
ments high in the atmosphere on weather balloons, air temperatures can
be measured without being affected by urbanization. Although data re-
corded this way show a consistent rise in global temperature, again such
measurements go back only a few decades.
Measurements of the level of greenhouse gases in the atmosphere are
also affected by urbanization. As a small town becomes a city, levels can be
expected to rise. However, recording stations located in regions far re-
moved from cities and factories also show an increase in the level of green-
house gases. One station on the island of Hawaii has shown a rise in the
amount of carbon dioxide present in the air since early 1958, with similar
reports coming from stations in Point Barrow, Alaska, and Antarctica.
Another important variable in looking at global warming is sea surface
temperature. Measurements can be skewed by local effects that have no
impact on the global climate. One method used to make detailed measure-
ments of seawater temperature is to broadcast a particular frequency of
noise through the water and measure it at distant locations. The speed and
frequency of the sound are affected by the temperature of the water. How-
ever, more accurate and more global data are becoming available through
observations made from satellites placed in geosynchronous orbits, which
can take a variety of detailed measurements of many conditions that are
factors in global warming. These data will help immensely in enabling sci-
entists to understand the global climate and the changes it is undergoing.
Christopher Keating
296
Global warming
See also: Biodiversity; Biomes: determinants; Biomes: types; Biosphere

concept; Ecosystems: definition and history; Ecosystems: studies; Geo-
chemical cycles; Greenhouse effect; Hydrologic cycle; Nutrient cycles;
Ozone depletion and ozone holes; Rain forests and the atmosphere;

Abrahamson, Dean Edwin, ed. The Challenge of Global Warming. Washing-
ton, D.C.: Island Press, 1989.
Berger, John J. Beating the Heat: Why and How We Must Combat Global
Warming. Berkeley, Calif.: Berkeley Hills Books, 2000.
Gates, David M. Climate Change and Its Biological Consequences. Sunderland,
Mass.: Sinauer Associates, 1993.
Houghton, John T. Global Warming: The Complete Briefing. 2d ed. New York:
Nance, John J. What Goes Up: The Global Assault on Our Atmosphere. New
York: William Morrow, 1991.
Rosenzweig, Cynthia. Climate Change and the Global Harvest: Potential Im-
pacts of the Greenhouse Effect on Agriculture. New York: Oxford Univer-
sity Press, 1998.
Sommerville, Richard C. J. The Forgiving Air: Understanding Environmental
Change. Berkeley: University of California Press, 1996.
Wyman, Richard L., ed. Global Climate Change and Life on Earth. New York:
Chapman and Hall, 1991.
297
GRASSLANDS AND PRAIRIES
Grasslands, including the prairies of North America, are a biome characterized by

the presence of low plants, mostly grasses, and are distinguished from woodlands,
deserts, and tundra. They support a great variety of plants and animals. At pres-
ent, the remaining grasslands provide grazing for livestock and wildlife.
G rasslands occupied vast areas of the world more than ten thousand
years ago, before the development of agriculture, industrialization,
and the subsequent explosive growth of the human population. They are
characterized by the presence of low plants (mostly grasses), experience
sparse to moderate rainfall, and are found in both temperate and tropical
climatic zones. The main grasslands of the planet include the prairies of
North America, the pampas of South America, the steppes of Eurasia, and
the savannas of Africa.
Grasslands are intermediate between deserts and woodlands in terms
of precipitation and biomass. The warmer tropical savannas average 60 to
150 centimeters (25 to 60 inches) of rain. The temperate grasslands range
between twenty-five to seventy-five centimeters (ten to thirty inches) of
precipitation, some of which may be in the form of snow. The biomass of
grasslands, predominantly grasses, is quantitatively intermediate between
that of deserts and woodlands, which produce 10 to 15 percent and 200 to
300 percent, respectively, of the amount of plant material. It should be rec-
ognized that the grassland biomes can be subdivided in terms of climate,
plant species, and animal species. It should also be noted that grasslands
do not always shift abruptly to deserts or woodlands, leading to grada-
tions between them. In addition, grasslands do have scattered trees, often
along streams or lakes, and low-lying brush.
Grasses have extensive root systems and the ability to become dormant.
These permit them to survive low rainfall, including periodic droughts, or
the winter cold typical of temperate regions. Furthermore, grasslands have
always been subjected to periodic fires, but the deep root systems of grass-
land plants also permit them to regrow after fire. Grasses coevolved over
millions of years with the grazing animals that depend on them for food.
Ten thousand years ago, wild ancestors of cattle and horses, as well as ante-
lope and deer, were on the Eurasian steppes; bison and pronghorn pros-
pered on the North American prairies; wildebeest, gazelle, zebra, and buf-
falo dominated African savannas; and the kangaroo was the predominant
298
Grasslands and prairies
grazer in Australia. Grazing is a symbiotic relationship, whereby animals

gain their nourishment from plants, which in turn benefit from the activity.
It removes vegetative matter, which is necessary in order for grasses to
grow, facilitates seed dispersal, and disrupts mature plants, permitting
young plants to take hold. Urine and feces from grazing animals recycle
nutrients to the plants. The grassland ecosystem also includes other ani-
mals, including worms, insects, birds, reptiles, rodents, and predators. The
grasses, grazing animals, and grassland carnivores, such as wolves or large
cat species, constitute a food chain.
Humans have been an increasing presence in grassland areas, where
more than 90 percent of contemporary crop production now occurs and
much urbanization and industrialization have taken place. Remaining
grassland areas are not used for crops, habitation, or industry because of
inadequate water supplies or unsuitable terrain but instead are used for
grazing domesticated or wild herbivores. In addition, many woodland ar-
eas around the world have been cleared and converted to grasslands for
crops, livestock, living, or working.
The Prairies of North America

Originally stretching east from the Rocky Mountains to Indiana and Ohio,
and from Alberta, Canada to Texas, the prairies were the major grassland
of North America. The short-grass prairie extended about 200 miles (300
kilometers) east of the mountains, and the long-grass prairie bordered the
deciduous forest along the eastern edge, while the mixed-grass prairie was
between the two. Going from west to east, the amount of precipitation in-
creases, causing changes in plant populations. The short-grass prairie re-
ceives only about 25 centimeters (10 inches) of precipitation each year,
mostly as summer rain, and, as its name suggests, has short grass, less than
60 centimeters (2 feet) tall. Today, it is used primarily for grazing because
the soil is shallow and unsuited for farming without irrigation. The mixed-
grass prairie receives moderate precipitation, ranging from 35 to 60 centi-
meters (14 to 24 inches) and has medium-height grasses, ranging from 60
to 120 centimeters (2 to 4 feet) tall. Much of it is now used for growing
wheat. The tall-grass prairie receives more than 60 centimeters (24 inches)
of precipitation, mostly in the summer, and had grasses that grow to over
150 centimeters (5 feet) tall. It has rich soil and has been mostly converted
to very productive cropland, primarily for corn and soybeans. The prairies
experience very cold winters (down to -45 degrees Celsius, -50 degrees
Fahrenheit) and very hot summers (up to 45 degrees Celsius, 110 degrees
Fahrenheit). They are often windy and experience severe storms, blizzards
in winter, thunderstorms and tornadoes in summer.
299
Grasses have extensive root systems and the ability to become dormant, permitting
them to survive the low to moderate rainfall that characterizes regions on the dry
sides of mountain ranges. The grassland ecosystem includes native grazers such as
bison or, more often today, domesticated grazers such as cattle, as well as worms,
insects, birds, reptiles, rodents, and predators such as wolves or large cats.
(PhotoDisc)
Like other biomes, the prairies have a characteristic assortment of ani-

mals, herbivores that eat the plants and carnivores that prey on the herbi-
vores. Before 1500 c.e., two ruminants, the bison (commonly but inaccu-
rately called buffalo) and the pronghorn (not a true antelope), were the
major grazers on the prairies. The prairie dog, a herbivorous rodent that
burrows, lived in large communities on the prairies. The major predators
were the wolf and coyote for bison and pronghorn, and the black-footed
ferret and fox for prairie dogs. A variety of birds (herbivorous and carnivo-
rous), reptiles, and insects also made their home on the prairie.
Overgrazing Grasslands
While grazing is of mutual benefit to plant and animal, overgrazing is ulti-
mately detrimental to both the plant and animal populations, as well as the
environment. Continued heavy grazing leads to deleterious consequences.
Removal of leaf tips, even repeated, will not affect regeneration of grasses
provided that the basal zone of the plant remains intact. While the upper
half of the grass shoot can generally be eaten without deleterious conse-
quences, ingesting the lower half, which sustains the roots and fuels
regrowth, will eventually kill the plants. Overgrazing leads to denuding
300
the land, to invasion by less nutritious plant species, to erosion due to de-
creased absorption of rainwater, and to starvation of the animal species.
Because the loss of plant cover changes the reflectance of the land, climate
changes can follow and make it virtually impossible for plants to return,
with desertification an ultimate consequence. It is not just the number of
animals, but the timing of the grazing that can be detrimental. Grasses re-
quire time to regenerate, and continuous grazing will inevitably kill them.
Consumption too early in the spring can stunt their development.
Semiarid regions are particularly prone to overgrazing because of low
and often unpredictable rainfall; regrettably, these are the areas of the
world where much grazing has been relegated, because the moister grass-
land areas have been converted to cropland. Overgrazing has contributed
to environmental devastation worldwide. Excessive grazing by cattle,
sheep, goats, and camels is partly responsible for the desert of the Middle
East, ironically the site of domestication for many animals and plants. Un-
controlled livestock grazing in the late 1800’s and early 1900’s negatively
affected many areas of the American West, where sagebrush and juniper
trees have invaded the grasslands. Livestock overgrazing has similarly
devastated areas of Africa and Asia. In the early twenty-first century, feral
horses in the American West and the Australian outback are damaging
those environments. Overgrazing by wildlife can also be deleterious. The
1924 Kaibab Plateau deer disaster in the Grand Canyon National Park and
Game Preserve is one such example, where removal of natural predators
led to overpopulation, overgrazing, starvation, and large die-offs.
Riparian zones, the strips of land on either side of a river or stream, are
particularly susceptible to overgrazing. Because animals naturally congre-
gate in these areas with water, lush vegetation, and shade, they can seri-
ously damage them by preventing grasses from regrowing and young
trees from taking root, as well as trampling and compacting the soil and
fouling the water course. The ecosystem can be devastated, threatening
survival of plant and animal species and leading to serious erosion. While
herding and fencing can be used to control animals in these areas, a less ex-
pensive method is to disperse the location of water supplies and salt blocks
to encourage movement away from rivers or streams. Grassland animals
crave salt, and if deprived of it, will seek it out.
Grassland Management
Grassland areas need not deteriorate if properly managed, whether for
livestock, wildlife, or both. Managing grasslands involves controlling the
number of animals and enhancing their habitat. Carrying capacity, which
is the number of healthy animals that can be grazed indefinitely on a given
301
unit of land, must not be exceeded. Because of year-to-year changes in

weather conditions and hence food availability, determining carrying ca-
pacity is not simple; worst-case estimates are preferred in order to mini-
mize the chances of exceeding it. The goal should be a healthy grassland
achieved by optimizing, not maximizing, the number of animals. For pri-
vate land, optimizing livestock numbers is in the long-term interest of the
landowner, although not always seen as such. For land that is publicly
held, managed in common, or with unclear or disputed ownership, re-
stricting animals to the optimum level is particularly difficult to achieve.
Personal short-term benefit often leads to long-term disaster, described as
the “tragedy of the commons” by biologist Garrett Hardin.
Appropriate management of grasslands involves controlling animal
numbers and enhancing grassland plants. Restricting cattle and sheep is
physically easy through herding and fencing, although it can be politically
difficult and expensive. Much more problematic is controlling charismatic
feral animals, such as horses, or wildlife, when natural predators have
been eliminated and hunting is severely restricted. As for habitat improve-
ment, the use of chemical, fire, mechanical, and biological approaches can
increase carrying capacity for either domesticated or wild herbivores. Re-
moving woody vegetation by burning or mechanical means will increase
grass cover, fertilizing can stimulate grass growth, and reseeding with de-
sirable species can enhance the habitat. Plants native to a particular region
can be best for preserving that environment. Effective grassland manage-
ment requires matching animals with the grasses on which they graze.
James L. Robinson
See also: Biomes: determinants; Biomes: types; Chaparral; Deserts; For-

ests; Habitats and biomes; Lakes and limnology; Marine biomes; Mediter-
ranean scrub; Mountain ecosystems; Old-growth forests; Rain forests; Rain
forests and the atmosphere; Rangeland; Reefs; Savannas and deciduous
tropical forests; Taiga; Tundra and high-altitude biomes; Wetlands.

Brown, Lauren. The Audubon Society Nature Guides: Grasslands. New York:
Knopf, 1985.
Collinson, Alan. Grasslands. New York: Dillon Press, 1992.
Demarais, Stephen, and Paul R. Krausman, eds. Ecology and Management of
Large Mammals in North America. Upper Saddle River, N.J.: Prentice
Hall, 2000.
Humphreys, L. R. The Evolving Science of Grassland Improvement. New York:
302
Joern, Anthony, and Kathleen H. Keeler, eds. The Changing Prairie. New
Pearson, C. J., and R. L. Ison. Agronomy of Grassland Systems. 2d ed. New
Sampson, Fred B., and Fritz L. Knopf, eds. Prairie Conservation. Washing-
ton, D.C.: Island Press, 1996.
Steele, Philip. Grasslands. Minneapolis: Carolrhoda Books, 1997.
303
GRAZING AND OVERGRAZING
Types of ecology: Agricultural ecology; Restoration and conservation

ecology
Animals that eat grass, or graze, can actually help the earth produce richer land
cover and soil. When the land suffers ill effects because of too much grazing, over-
grazing has occurred.
T he effects of overgrazing occur where there are more grazing animals

than the land and vegetation can support. Overgrazing has negatively
affected regions of the United States, primarily in the Southwest. Areas
that have been severely damaged by overgrazing typically show declining
or endangered plant and animal species.
Herbivores are animals that feed on plant material, and grazers are her-
bivores that feed specifically on grass. Examples are horses, cows, ante-
lope, rabbits, and grasshoppers. Overgrazing occurs when grazer popula-
tions exceed the carrying capacity of a specified area (the number of
individual organisms the resources of a given area can support). In over-
grazing conditions, there is insufficient food to support the animal popula-
tion in question. Depending on the grazer’s strategy, emigration or starva-
tion will follow. Grasslands can handle, and even benefit from, normal
grazing; only overgrazing adversely affects them.
Grasses’ Defenses Against Grazing

Grasslands and grazers coevolved, so grasses can withstand grazing
within the ecosystem’s carrying capacity. All plants have a site of new cell
growth called the meristem, where growth in height and girth occurs.
Most plants have the meristem at the very top of the plant (the apical
meristem). If a plant’s apical meristem is removed, the plant dies.
If grasses had an apical meristem, grazers—and lawn mowers—would
kill grasses. Grasses survive mowing and grazing because the meristem is
located at the junction of the shoot and root, close to the ground. With the
exception of sheep, grazers in North America do not disturb the meristem,
and sheep do so only during overgrazing conditions. At proper levels of
grazing, grazing actually stimulates grass to grow in height in an attempt
to produce a flowering head for reproduction. Grazing also stimulates
grass growth by removing older plant tissue at the top that is functioning
at a lower photosynthetic rate.
304
Grazing and overgrazing
Grazers
Mammalian grazers have high, crowned teeth with a great area for grind-
ing to facilitate opening of plants’ cell walls as a means to release nutrients.
The cell wall is composed of cellulose, which is very difficult for grazers to
digest. Two major digestive systems of grazing strategies have evolved to
accommodate grazing. Ruminants, such as cows and sheep, evolved stom-
achs with four chambers to allow regurgitation in order to chew food twice
to maximize cellulose breakdown. Intestinal bacteria digest the cellulose,
releasing fatty acids that nourish the ruminants. Other grazers, such as
rabbits and horses, house bacteria in the cecum, a pouch at the junction of
the small and large intestines. These bacteria ferment the plant material in-
gested. The fermented products of the bacteria nourish these grazers.
Impacts in the Southwest

As previously mentioned, in the United States the negative effects of over-
grazing are most intense in the Southwest. Some ecologists believe that one
significant factor was the pattern of early European colonization of the
area. Missions were abundant in the Southwest, and the missions owned
Grazers such as sheep form a symbiotic relationship with the grasses, getting their
nourishment from these plants and in turn facilitating growth of new grass by re-
moving excess vegetative matter, dispersing seed through their droppings, and re-
cycling nutrients via urine and feces. Overgrazing occurs when the number of
grazers exceeds the carrying capacity of the grassland. (PhotoDisc)
305
cattle that were rarely slaughtered, except on big feast days. Because Cath-
olic missionaries received some financial support from their religious or-
ders in Europe, mission cattle were not restrained as strictly as were those
owned by cattle ranchers, whose sole livelihood came from raising and
selling cattle. Mission cattle roamed greater distances and began the pat-
tern of overgrazing in the Southwest. The impact of overgrazing was par-
ticularly intense because much of the Southwest has desertlike conditions.
Extreme environmental conditions result in particularly fragile ecosys-
tems. Hence the Southwest was, and is, vulnerable to the effects of over-
grazing.
Another possible—though disputed—contribution to overgrazing may
stem from the fact that much of the land in the Southwest is public land un-
der jurisdiction of the Bureau of Land Management. This federal agency
leases land to private concerns for the purpose of grazing cattle or sheep.
Some observers feel that the bureau has a conflict of interest in that its pri-
mary source of income is money obtained from leasing public land under
its jurisdiction. They suspect that the bureau has granted, and fear that it
may continue to grant, grazing leases in regions threatened with or suffer-
ing from overgrazing.
Effects of Overgrazing
Overgrazing can lead to a number of ecological problems. Depletion of
land cover leads to soil erosion and can ultimately cause desertification.
Other possible results are the endangering of some species of grass and the
creation of monocultures in regions where certain species have been re-
moved. Desertification is the intensification and expansion of deserts at the
expense of neighboring grasslands. When overgrazing occurs along desert
perimeters, plant removal leads to decreased shading. Decreased shading
increases the local air temperature. When the temperature increases, the air
may no longer cool enough to release moisture in the form of dew. Dew is
the primary source of precipitation in deserts, so without it, desert condi-
tions intensify. Even a slight decrease in desert precipitation is serious. The
result is hotter and drier conditions, which lead to further plant loss and
potentially to monocultures.
Overgrazing of grasslands, combined with the existence of nonnative
species in an ecosystem, can result in the endangerment of species of native
grasses. At one time, cattle in the Southwest fed exclusively on native
grasses. Then nonnative plant species arrived in the New World in the guts
of cows shipped from Europe. They began to compete with the native
grasses. European grass species have seeds with prickles and burs; south-
western native grasses do not, making them softer and more desirable to
306
the cattle. Hence European grasses experienced little, if any, grazing, while
the much more palatable southwestern native grasses were grazed to the
point of overgrazing. The result was drastic decline or loss of native grass-
land species. In such cases animals dependent on native grassland species
must emigrate or risk extinction. For example, many ecologists conjecture
that the Coachella Valley kit fox in California is threatened because of the
loss of grassland habitat upon which it is dependent.
Solutions
Desertification is usually considered irreversible, but the elimination of
grazing along desert perimeters can help to prevent further desertification.
One kind of attempt to reestablish native grass species involves controlled-
burn programs. Nonnative grassland species do not appear to be as fire-
resistant as native grass species. Controlled burn programs are therefore
being used in some overgrazed grassland areas to try to eliminate nonna-
tives and reestablish native grass species. If successful, such programs will
improve the health of the ecosystem.
Jessica O. Ellison
See also: Biopesticides; Desertification; Erosion and erosion control; Forest

fires; Forest management; Forests; Genetically modified foods; Grazing
and overgrazing; Integrated pest management; Multiple-use approach;
Pesticides; Rangeland; Slash-and-burn agriculture; Soil; Soil contamina-
tion; Wildlife management.

Hodgson, J., and A. W. Illius, eds. The Ecology and Management of Grazing
Systems. Wallingford, Oxon, England: CAB International, 1996.
McBrien, Heather, et al. “A Case of Insect Grazing Affecting Plant Succes-
sion.” Ecology 64, no. 5 (1983).
Sousa, Wayne P. “The Role of Disturbance in Natural Communities.” An-
nual Review Ecological Systems 15, 1984.
_______. “Some Basic Principles of Grassland Fire Management.” Environ-
mental Management 3, no. 1 (1979).
WallisDeVries, Michiel F., Jan P. Bakker, Sipke E. Van Wieren, eds. Grazing
and Conservation Management. Boston: Kluwer Academic, 1998.
307
GREENHOUSE EFFECT
Type of ecology: Global ecology
The greenhouse effect is a natural process of atmospheric warming in which solar en-
ergy that has been absorbed by the earth’s surface is reradiated and then absorbed by
particular atmospheric gases, primarily carbon dioxide and water vapor. Without
this warming process, the atmosphere would be too cold to support life. Since 1880,
however, the surface atmospheric temperature has been rising, paralleling a rise in
the concentration of carbon dioxide and other gases produced by human activities.
S ince 1880, carbon dioxide, along with several other gases—chloro-

fluorocarbons (CFCs), methane, hydrofluorocarbons (HFCs), per-
fluourocarbons (PFCs), sulfur hexafluoride, and nitrous oxide—have been
increasing in concentration and have been identified as likely contributors
to a rise in global surface temperature. These gases are called greenhouse
gases. The temperature increase may lead to drastic changes in climate and
food production as well as widespread coastal flooding. As a result, many
scientists, organizations, and governments have called for curbs on the
production of greenhouse gases. Since the predictions are not definite,
however, debate continues about the costs of reducing the production of
these gases without being sure of the benefits.
Global Warming
The greenhouse effect occurs because the gases in the atmosphere are able
to absorb only particular wavelengths of energy. The atmosphere is largely
transparent to short-wave solar radiation, so sunlight basically passes
through the atmosphere to the earth’s surface. Some is reflected or ab-
sorbed by clouds, some is reflected from the earth’s surface, and some is
absorbed by dust or the earth’s surface. Only small amounts are actually
absorbed by the atmosphere. Therefore, sunlight contributes very little to
the direct heating of the atmosphere. On the other hand, the greenhouse
gases are able to absorb long-wave, or infrared, radiation from the earth,
thereby heating the earth’s atmosphere.
Discussion of the greenhouse effect has been confused by terms that are
imprecise and even inaccurate. For example, the atmosphere was believed
to operate in a manner similar to a greenhouse, whose glass would let visi-
ble solar energy in but would also be a barrier preventing the heat energy
from escaping. In actuality, the reason that the air remains warmer inside a
greenhouse is probably because the glass prevents the warm air from mix-
ing with the cooler outside air. Therefore the greenhouse effect could be
308
Greenhouse effect
more accurately called the “atmospheric effect,” but the term greenhouse
effect continues to be used.
Even though the greenhouse effect is necessary for life on earth, the term
gained harmful connotations with the discovery of apparently increasing at-
mospheric temperatures and growing concentrations of greenhouse gases.
The concern, however, is not with the greenhouse effect itself but rather
with the intensification or enhancement of the greenhouse effect, presum-
ably caused by increases in the level of gases in the atmosphere resulting
from human activity, especially industrialization. Thus the term global
warming is a more precise description of this presumed phenomenon.
The Greenhouse Effect
Sun
Earth
Atmosphere
Clouds and atmospheric gases such as water vapor, carbon dioxide, meth-
ane, and nitrous oxide absorb part of the infrared radiation emitted by the
earth’s surface and reradiate part of it back to the earth. This process effectively
reduces the amount of energy escaping into space and is popularly called the
“greenhouse effect” because of its role in warming the lower atmosphere.
The greenhouse effect has drawn worldwide attention because increasing
concentrations of carbon dioxide from the burning of fossil fuels may result
in a global warming of the atmosphere. Scientists know that the greenhouse
analogy is incorrect. A greenhouse traps warm air within a glass building
where it cannot mix with cooler air outside. In a real greenhouse, the trap-
ping of air is more important in maintaining the temperature than is the trap-
ping of infrared energy. In the atmosphere, air is free to mix and move about.
309
Greenhouse effect
Human Contributions
A variety of human activities appear to have contributed to global warm-
ing. Large areas of natural vegetation and forests have been cleared for ag-
riculture. The crops may not be as efficient in absorbing carbon dioxide as
the natural vegetation they replaced. Increased numbers of livestock have
led to growing levels of methane. Several gases that appear to be intensify-
ing global warming, including CFCs and nitrous oxides, also appear to be
involved with ozone depletion. Stratospheric ozone shields the earth from
solar ultraviolet (short-wave) radiation; therefore, if the concentration of
these ozone-depleting gases continues to increase and the ozone shield is
depleted, the amount of solar radiation reaching the earth’s surface should
increase. Thus, more solar energy would be intercepted by the earth’s sur-
face to be reradiated as long-wave radiation, which would presumably in-
crease the temperature of the atmosphere.
However, whether there is a direct cause-and-effect relationship be-
tween increases in carbon dioxide and the other gases and surface temper-
ature may be impossible to determine because the atmosphere’s tempera-
ture has fluctuated widely over millions of years. Over the past 800,000
years, the earth has had several long periods of cold temperatures—during
which thick ice sheets covered large portions of the earth—interspersed
with shorter warm periods. Since the most recent retreat of the glaciers
around 10,000 years ago, the earth has been relatively warm.
Problems of Prediction
How much the temperature of the earth might rise is not clear. So far, the
temperature increase of around 1 degree Fahrenheit is within the range of
normal (historic) trends. The possibility of global warming became a seri-
ous issue during the late twentieth century because the decades of the
1980’s and the 1990’s included some of the hottest years recorded for more
than a century. On the other hand, warming has not been consistent since
1880, and for many years cooling occurred. The cooling might have re-
sulted from the increase of another product of fossil fuel combustion, sul-
fur dioxide aerosols, which reflect sunlight, thus lessening the amount of
solar energy entering the atmosphere. Similarly, in the early 1990’s temper-
atures declined, perhaps because of ash and sulfur dioxide produced by
large volcanic explosions during that period. In the late 1990’s and early
2000’s temperatures appeared to be rising again, thus indicating that prod-
ucts of volcanic explosions may have masked the process of global warm-
ing. The United States Environmental Protection Agency (EPA) states that
the earth’s average temperature will probably continue to increase because
the greenhouse gases stay in the atmosphere longer than the aerosols.
310
Greenhouse effect
Proper analysis of global warming is dependent on the collection of ac-

curate temperature records from many locations around the world and
over many years. Because human error is always possible, “official” tem-
perature data may not be accurate. This possibility of inaccuracy compro-
mises examination of past trends and predictions for the future. However,
the use of satellites to monitor temperatures has probably increased the re-
liability of the data.
Predictions for the future are hampered in various ways, including lack
of knowledge about all the components affecting atmospheric tempera-
ture. Therefore, computer programs cannot be sufficiently precise to make
accurate predictions. A prime example is the relationship between ocean
temperature and the atmosphere. As the temperature of the atmosphere
increases, the oceans would absorb much of that heat. Therefore, the atmo-
sphere might not warm as quickly as predicted. However, the carbon diox-
ide absorption capacity of oceans declines with temperature. Therefore,
the oceans would be unable to absorb as much carbon dioxide as before,
but exactly how much is unknown. Increased ocean temperatures might
also lead to more plant growth, including phytoplankton. These plants
would probably absorb carbon dioxide through photosynthesis. A warmer
atmosphere could hold more water vapor, resulting in the potential for
more clouds and more precipitation. Whether that precipitation would fall
as snow or rain and where it would fall could also affect air temperatures.
Air temperature could lower as more clouds might reflect more sunlight,
or more clouds might absorb more infrared radiation.
To complicate matters, any change in temperature would probably not
be uniform over the globe. Because land heats up more quickly than water,
the Northern Hemisphere, with its much larger landmasses, would proba-
bly have greater temperature increases than the Southern Hemisphere.
Similarly, ocean currents might change in both direction and temperature.
These changes would affect air temperatures as well. In reflection of these
complications, computer models of temperature change range widely in
their estimates. Predicted increases range from 1.5 to 11 degrees Celsius (3
to 20 degrees Fahrenheit) over the early decades of the twenty-first cen-
tury.
Mitigation Attempts
International conferences have been held, and international organizations
have been established to research and minimize potential detriments of
global warming. In 1988 the United Nations Environment Programme and
the World Meteorological Organization established the International Panel
on Climate Change (IPCC). The IPCC has conducted much research on cli-
311
Greenhouse effect
mate change and is now considered an official advisory body on the cli-
mate change issue. In June, 1992, the United Nations Conference on Envi-
ronment and Development, or Earth Summit, was held in Brazil.
Participants devised the Framework Convention on Climate Change and
considered the landmark international treaty. It required signatories to re-
duce and monitor their greenhouse gas emissions.
A more advanced agreement, the Kyoto Accords, was developed in De-
cember, 1997, by the United Nations Framework Convention on Climate
Change. It set binding emission levels for all six greenhouse gases over a
five-year period for the developed world. Developing countries do not
have any emission targets. It also allows afforestation to be used to offset
emissions targets. The Kyoto agreement includes the economic incentive
of trading emissions targets. Some countries, because they have met their
targets, would have excess permits, which they might be willing to sell to
other countries that have not met their targets.
Margaret F. Boorstein
See also: Biodiversity; Biomes: determinants; Biomes: types; Biosphere

concept; Geochemical cycles; Global warming; Hydrologic cycle; Ozone
depletion and ozone holes; Rain forests and the atmosphere.

Berger, John J. Beating the Heat: Why and How We Must Combat Global
Warming. Berkeley, Calif.: Berkeley Hills Books, 2000.
Flavin, Christopher, Odil Tunali, and Jane A. Peterson, et al., eds. Climate of
Hope: New Strategies for Stabilizing the World’s Atmosphere. Washington,
D.C.: Worldwatch Institute, 1996.
Graedel, Thomas E. Atmosphere, Climate, and Change. New York: Scientific
American Library/W. H. Freeman, 1997.
Houghton, John T. Global Warming: The Complete Briefing. 2d ed. New York:
Kondratsev, Kirill, and Arthur P. Cracknell. Observing Global Climate Change.
London: Taylor and Francis, 1998.
McKibben, Bill. The End of Nature. 10th anniversary ed. New York: Anchor
Books, 1999.
Rosenzweig, Cynthia. Climate Change and the Global Harvest: Potential Im-
pacts of the Greenhouse Effect on Agriculture. New York: Oxford Univer-
sity Press, 1998.
Sommerville, Richard C. J. The Forgiving Air: Understanding Environmental
Change. Berkeley: University of California Press, 1996.
312
HABITATS AND BIOMES
The biosphere is the sum total of all habitats on earth that can be occupied by living
organisms. Descriptive and experimental studies of habitat components allow sci-
entists to predict how various organisms will respond to changes in their environ-
ment, whether caused by humans or nature.
L ife in the form of individual organisms composed of one or more living

cells is found in a vast array of different places on earth, each with its
own distinctive types of organisms. Life on earth has been classified by sci-
entists into units called species, whose individuals appear similar, have the
same role in the environment, and breed only among themselves. The
space in which each species lives is called its habitat.
Habitats, Communities, Ecosystems, and Biomes

The term “habitat” can refer to specific places with varying degrees of ac-
curacy. For example, rainbow trout can be found in North America from
Canada to Mexico, but more specifically they are found in freshwater
streams and lakes, with an average temperature below 70 degrees Fahren-
heit and a large oxygen supply. The former example describes the macro-
habitat of the rainbow trout, which is a broad, easily recognized area. The
latter example describes its microhabitat—the specific part of its macro-
habitat in which it is found. Similarly, the macrohabitat of one species can
refer to small or large areas of its habitat. The macrohabitat of rainbow
trout may refer to the habitat of a local population, the entire range of the
species, or (most often) an area intermediate to those extremes. While
“habitat,” therefore, refers to the place an organism lives, it is not a precise
term unless a well-defined microhabitat is intended.
The total population of each species has one or more local populations,
which are all the individuals in a specific geographic area that share a com-
mon gene pool; that is, they commonly interbreed. For example, rainbow
trout of two adjacent states will not normally interbreed unless they are
part of local populations that are very close to each other. The entire geo-
graphic distribution of a species, its range, may be composed of many local
populations.
On a larger organizational scale, there is more than only one local popu-
lation of one species in any habitat. Indeed, it is natural and necessary for
many species to live together in an area, each with its own micro- and
313
Habitats and biomes
macrohabitat. The habitat of each local population of each species overlaps

the habitat of many others. This collective association of populations in one
general area is termed a community, which may consist of thousands of
species of animals, plants, fungi, bacteria, and other one-celled organisms.
Groups of communities that are relatively self-sufficient in terms of
both recycling nutrients and the flow of energy among them are called eco-
systems. An example of an ecosystem could be a broad region of forest
community interspersed with meadows and stream communities that
share a common geographic area. Some ecosystems are widely distributed
across the surface of the earth and are easily recognizable as similar ecosys-
tems known as biomes—deserts, for example. Biomes are usually named
for the dominant plant types, which have very similar shapes and macro-
habitats. Thus, similar types of organisms inhabit them, though not neces-
sarily ones of the same species. These biomes are easily mapped on the
continental scale and represent a broad approach to the distribution of or-
ganisms on the face of the earth. One of the more consistent biomes is the
northern coniferous forest, which stretches across Canada and northern
Eurasia in a latitudinal belt. Here are found needle-leafed evergreen coni-
fer trees adapted to dry, cold, windy conditions in which the soil is frozen
during the long winter.
North American Biomes

The biomes of North America, from north to south, are the polar ice cap,
the Arctic tundra, the northern coniferous forest; then, at similar middle
latitudes, eastern deciduous forest, prairie grassland, or desert; and last,
subtropical rain forest near the equator. Complicating factors that deter-
mine the actual distribution of the biomes are altitude, annual rainfall, to-
pography, and major weather patterns. These latter factors, which influ-
ence the survival of the living, or biotic, parts of the biome, are called
abiotic factors. These are the physical components of the environment for a
community of organisms.
The polar ice cap is a hostile place with little evidence of life on the sur-
face except for polar bears and sea mammals that depend on marine ani-
mals for food. A distinctive characteristic of the Arctic tundra, just south of
the polar ice cap, is its flat topography and permafrost, or permanently
frozen soil. Only the top meter or so thaws during the brief Arctic summer
to support low-growing mosses, grasses, and the dominant lichens known
as reindeer moss. Well-known animals found there are the caribou, musk-
ox, lemming, snowy owl, and Arctic fox.
The northern coniferous forest is dominated by tall conifer trees. Famil-
iar animals include the snowshoe hare, lynx, and porcupine. This biome
314
Habitats and biomes
stretches east to west across Canada and south into the Great Lakes region
of the United States. It is also found at the higher elevations of the Rocky
Mountains and the western coastal mountain ranges. Its upper elevation
limit is the “treeline,” above which only low-growing grasses and herba-
ceous plants grow in an alpine tundra community similar to the Arctic tun-
dra. In mountain ranges, the change in biomes with altitude mimics the
biome changes with increasing latitude, with tundra being the highest or
northernmost.
Approximately the eastern half of the United States was once covered
with the eastern deciduous forest biome, named for the dominant broad-
leaved trees that shed their leaves in the fall. This biome receives more than
75 centimeters (about 30 inches) of rainfall each year and has a rich diver-
sity of bird species, such as the familiar warblers, chickadees, nuthatches,
and woodpeckers. Familiar mammals include the white-tailed deer, cot-
tontail rabbit, and wild turkey. The Great Plains, between the Mississippi
River and the Rocky Mountains, receives 25 to 75 centimeters (about 10 to
30 inches) of rain annually to support an open grassland biome often called
the prairie. The many grass species that dominate this biome once sup-
ported vast herds of bison and, in the western parts, pronghorn antelope.
Seasonal drought and periodic fires are common features of grasslands.
The land between the Rockies and the western coastal mountain ranges
is a cold type of desert biome; three types of hotter deserts are found from
western Texas west to California and south into Mexico. Deserts receive
fewer than 25 centimeters (10 inches) of rainfall annually. The hot deserts
are dominated by many cactus species and short, thorny shrubs and trees,
whereas sagebrush, grass, and small conifer trees dominate the cold
desert. These deserts have many lizard and snake species, including poi-
sonous rattlesnakes and the Gila monster. The animals often have noctur-
nal habits to avoid the hot, dry daytime.
Southern Mexico and the Yucatan Peninsula are covered by evergreen,
broad-leaved trees in the tropical rain-forest biome, which receives more
than 200 centimeters (almost 80 inches) of rain per year. Many tree-dwell-
ing animals, such as howler monkeys and tree frogs, spend most of their
lives in the tree canopy, seldom reaching the ground.
Aquatic biomes can be broadly categorized into freshwater, marine, and
estuarine biomes. Freshwater lakes, reservoirs, and other still-water envi-
ronments are called lentic, in contrast to lotic, or running-water, environ-
ments. Lentic communities are often dominated by planktonic organisms,
small, drifting (often transparent) microscopic algae, and the small ani-
mals that feed on them. These, in turn, support larger invertebrates and
fish. Lotic environments depend more on algae that are attached to the bot-
315
Habitats and biomes
toms of streams, but they support equally diverse animal communities.

The marine biome is separated into coastal and pelagic, or open-water, en-
vironments, which have plant and animal communities somewhat similar
to lotic and lentic freshwater environments, respectively. The estuarine
biome is a mixing zone where rivers empty into the ocean. These areas
have a diverse assemblage of freshwater and marine organisms.
The Biosphere
All the biomes together, both terrestrial and aquatic, constitute the bio-
sphere, which by definition is all the places on earth where life is found.
Organisms that live in a biome must interact with one another and must
successfully overcome and exploit their abiotic environment. The severity
and moderation of the abiotic environments determine whether life can ex-
ist in that microhabitat. Such things as minimum and maximum daily and
annual temperature, humidity, solar radiation, rainfall, and wind speed di-
rectly affect which types of organisms are able to survive. Amazingly, few
places on earth are so hostile that no life exists there. An example would be
the boiling geyser pools at Yellowstone National Park, but even there, as
the water temperature cools at the edges to about 75 degrees Celsius, bacte-
rial colonies begin to appear. There is abundant life in the top meter or two
of soil, with plant roots penetrating to twenty-two meters or more in ex-
treme cases. Similarly, the mud and sand bottoms of lakes and oceans con-
tain a rich diversity of life. Birds, bats, and insects exploit the airspace
above land and sea up to a height of about 1,200 meters (nearly 4,000 feet),
with bacterial and fungal spores being found much higher. Thus, the bio-
sphere generally extends about 10 to 15 meters (33 to 50 feet) below the sur-
face of the earth and about 1,200 meters above it. Beyond that, conditions
are too hostile. A common analogy is that if the earth were a basketball, the
biosphere would constitute only the thin outer layer.
Studying Habitats and Biomes

Abiotic habitat requirements for a local population or even for an entire
species can be determined in the laboratory by testing its range of tolerance
for each factor. For example, temperature can be regulated in a laboratory
experiment to determine the minimum and maximum survival tempera-
tures as well as an optimal range. The same can be done with humidity,
light, shelter, and substrate type: “Substrate preference” refers to the solid
or liquid matter in which an organism grows and/or moves—for example,
soil or rock. The combination of all ranges of tolerance for abiotic factors
should describe a population’s actual or potential microhabitat within a
community. Furthermore, laboratory experiments can theoretically indi-
316
Habitats and biomes
cate how much environmental change each population can tolerate before
it begins to migrate or die.
Methods to study the interaction of populations with one another or
even the interaction of individuals within one local population are much
more complicated and are difficult or impossible to bring into a laboratory
setting. These studies most often require collecting field data on distribu-
tion, abundance, food habits or nutrient requirements, reproduction and
death rates, and behavior in order to describe the relationships between in-
dividuals and populations within a community. Later stages of these field
investigations could involve experimental manipulations in which scien-
tists purposely change one factor, then observe the population or commu-
nity response. Often, natural events such as a fire, drought, or flood can
provide a disturbance in lieu of human manipulation.
There are obvious limits to how much scientists should tinker with the
biosphere merely to see how it works. Populations and even communities
in a local area can be manipulated and observed, but it is not practical or
advisable to manipulate whole ecosystems or biomes. To a limited extent,
scientists can document apparent changes caused by civilization, pollu-
tion, and long-term climatic changes. This information, along with popula-
tion- and community-level data, can be used to construct a mathematical
model of a population or community. The model can then be used to pre-
dict the changes that would happen if a certain event were to occur. These
predictions merely represent the “best guesses” of scientists, based on the
knowledge available. Population ecologists often construct reasonably ac-
curate population models that can predict population fluctuations based
on changes in food supply, abiotic factors, or habitat. As models begin to
encompass communities, ecosystems, and biomes, however, their knowl-
edge bases and predictive powers decline rapidly. Perhaps the most com-
plicating factor in building and testing these large-scale models is that nat-
ural changes seldom occur one at a time. Thus, scientists must attempt to
build cumulative-effect models that are capable of incorporating multiple
changes into a predicted outcome.
James F. Fowler
See also: Biomes: determinants; Biomes: types; Biosphere concept; Chap-

arral; Deserts; Ecosystems: definition and history; Ecosystems: studies;
Forests; Grasslands and prairies; Lakes and limnology; Marine biomes;
Mediterranean scrub; Mountain ecosystems; Old-growth forests; Rain for-
ests; Rain forests and the atmosphere; Rangeland; Reefs; Savannas and de-
ciduous tropical forests; Taiga; Tundra and high-altitude biomes; Wet-
lands.
317
Habitats and biomes

Allaby, Michael. Biomes of the World. 9 vols. Danbury, Conn.: Grolier Inter-
national, 1999.
Bradbury, Ian K. The Biosphere. New York: Belhaven Press, 1991.
Cox, George W. Conservation Biology: Concepts and Applications. 2d ed.
Dubuque, Iowa: Wm. C. Brown, 1997.
Hanks, Sharon La Bonde. Ecology and the Biosphere: Principles and Problems.
Delray Beach, Fla.: St. Lucie Press, 1996.
Luoma, Jon R. The Hidden Forest: The Biography of an Ecosystem. New York:
Henry Holt, 1999.
Miller, G. Tyler, Jr. Living in the Environment. 12th ed. Belmont, Calif.:
Thomson Learning, 2001.
Sutton, Ann, and Myron Sutton. Eastern Forests. New York: Alfred A.
Knopf, 1986.
Wetzel, Robert G. Limnology. 3d ed. San Diego: Academic Press, 2001.
318
HABITUATION AND
SENSITIZATION
Habituation is learning to ignore irrelevant stimuli that previously produced a re-

action. Results of habituation studies have been used to explain, predict, and con-
trol behavior of humans and other organisms.
H abituation is a simple form of nonassociative learning that has been

demonstrated in organisms as diverse as protozoans, insects, Nereis
(clam worms), birds, and humans. The habituated organism learns to ig-
nore irrelevant, repetitive stimuli which, prior to habituation, would have
produced a response. With each presentation of the habituating stimulus,
the responsiveness of the organism decreases toward the zero, nonresponse
level. If habituation training continues after the zero-response level, the
habituation period is prolonged. Habituation to a particular stimulus nat-
urally and gradually disappears unless the training continues. If training is
resumed after habituation has disappeared, habituation occurs more rap-
idly in the second training series than in the first. Habituation is important
for survival of the individual. Many stimuli are continuously impinging
upon it: Some are important, others are not. Important stimuli require an
immediate response, but those which result in neither punishment nor re-
ward may be safely ignored.
Stimulus and Response

When a new stimulus is presented (when a sudden change in the environ-
ment occurs), the organism—be it bird, beast, or human—exhibits the
“startle” or “orientation” response. In essence, it stops, looks, and listens. If
the stimulus is repeated and is followed by neither reward nor punish-
ment, the organism will pay less and less attention to it. When this hap-
pens, habituation has occurred, and the organism can now respond to and
deal with other stimuli. On the other hand, if, during habituation learning,
a painful consequence follows a previously nonconsequential stimulus,
the organism has been sensitized to that stimulus and will respond to it
even more strongly than it did before the learning sessions, whether they
are occurring in the laboratory or in the field.
Young birds must learn to tell the difference between and respond dif-
ferently to a falling leaf and a descending predator. A young predatory bird
319
Habituation and sensitization
must learn to ignore reactions of its prey which pose no danger, reactions
that the predator initially feared.
A theory known as the dual-process habituation-sensitization theory
was formulated in 1966 and revised in 1973. It establishes criteria for both
habituation and sensitization. Criteria for habituation (similar to those
proposed by E. N. Sokolov in 1960) are that habituation will develop rap-
idly; the frequency of stimulation determines the degree of habituation; if
stimulation stops for a period of time, habituation will disappear; the
stronger the stimulus, the slower the rate of habituation; the frequency of
stimulation is more important than the strength of the stimulus; rest peri-
ods between habituation series increase the degree of habituation; and the
organism will generalize and therefore exhibit habituation to an entire
class of similar stimuli. Stimulus generalization can be measured: If a dif-
ferent stimulus is used in the second habituation series, habituation occurs
more rapidly; it indicates generalization.
Sensitization
Sensitization, a very strong response to a very painful, injurious, or harm-
ful stimulus, is not limited to stimulus-response circuits but involves the
entire organism. After sensitization, the individual may respond more
strongly to the habituating stimulus than it did prior to the start of habitua-
tion training.
There are eight assumptions about sensitization in the dual-process the-
ory. Sensitization does not occur in stimulus-response circuits but involves
the entire organism. Sensitization increases during the early stages of ha-
bituation training but later decreases. The stronger the sensitizing stimulus
and the longer the exposure to it the greater the sensitization; weaker stim-
uli may fail to produce any sensitization. Even without any external inter-
vention, sensitization will decrease and disappear. Increasing the fre-
quency of sensitization stimulation causes a decrease in sensitization.
Sensitization will extend to similar stimuli. Dishabituation, the loss of ha-
bituation, is an example of sensitization. Sensitization may be time-related,
occurring only at certain times of the day or year.
According to the dual-process theory, the response of an organism to a
stimulus will be determined by the relative strengths of habituation and
sensitization. Charles Darwin, the father of evolution, observed and de-
scribed habituation, although he did not use the term. He noted that the
birds of the Galápagos Islands were not disturbed by the presence of the gi-
ant tortoises, Amblyrhynchus; they disregarded them just as the magpies in
England, which Darwin called “shy” birds, disregarded cows and horses
grazing nearby. Both the giant tortoises of the Galápagos Islands and the
320
grazing horses and cows of England were stimuli which, though present,
would not produce profit or loss for the birds; therefore, they could be ig-
nored.
The Neurology of Stimulus Response

Within the bodies of vertebrates is a part of the nervous system called the
reticular network or reticular activating system; it has been suggested that
the reticular network is largely responsible for habituation. It extends
from the medulla through the midbrain to the thalamus of the forebrain.
(The thalamus functions as the relay and integration center for impulses
to and from the cerebrum of the forebrain.) Because it is composed of a
huge number of interconnecting neurons and links all parts of the body,
the reticular network functions as an evaluating, coordinating, and alarm
center. It monitors incoming message impulses. Important ones are per-
mitted to continue to the cerebral cortex, the higher brain. Messages from
the cerebral cortex are coordinated and dispatched to the appropriate
areas.
During sleep, many neurons of the reticular network stop functioning.
Those that remain operational may inhibit response to unimportant stim-
uli (habituation) or cause hyperresponsiveness (sensitization). The cat
who is accustomed to the sound of kitchen cabinets opening will sleep
through a human’s dinner being prepared (habituation) but will charge
into the kitchen when the she hears the sound of the cat food container
opening (sensitization).
Researcher E. N. Sokolov concluded that the “orientation response”
(which can be equated with sensitization) and habituation are the result of
the functioning of the reticular network. According to Sokolov, habituation
results in the formation of models within the reticular activating system.
Incoming messages that match the model are disregarded by the organism,
but those that differ trigger alerting reactions throughout the body, thus
justifying the term “alerting system” as a synonym for the reticular net-
work. Habituation to a very strong stimulus would take a long time. Repe-
tition of this strong stimulus would cause an even stronger defensive reflex
and would require an even longer habituation period.
The Role of Neurotransmitters

Neurotransmitters are chemical messengers that enable nerve impulses to
be carried across the synapse, the narrow gap between neurons. They
transmit impulses from the presynaptic axon to the postsynaptic den-
drite(s). E. R. Kandell, in experiments with Aplysia (the sea hare, a large
mollusk), demonstrated that as a habituation training series continues,
321
smaller amounts of the neurotransmitter acetylcholine are released from

the axon of the presynaptic sensory neuron. On the other hand, after sensi-
tization, this neuron released larger amounts of acetylcholine because
of the presence of serotonin, a neurotransmitter secreted by a facilitory
interneuron. When a sensitizing stimulus is very strong, it usually gener-
ates an impulse within the control center—a ganglion, a neuron, or the
brain. The control center then transmits an impulse to a facilitory inter-
neuron, causing the facilitory interneuron to secrete serotonin.
Increased levels of acetylcholine secretion by the sensory neuron result
from two different stimuli: direct stimulation of the sensory neurons of the
siphon or serotonin from the facilitory interneuron. Facilitory interneurons
synapse with sensory neurons in the siphon. Serotonin discharged from
facilitory interneurons causes the sensory neurons to produce and secrete
more acetylcholine.
On the molecular level, the difference between habituation and adapta-
tion—the failure of the sensory neuron to respond—is very evident. The
habituated sensory neuron has a neurotransmitter in its axon but is unable
to secrete it and thereby enable the impulse to be transmitted across the
synapse. The adapted sensory neuron, by contrast, has exhausted its cur-
rent supply of neurotransmitter. Until new molecules of neurotransmitter
are synthesized within the sensory neuron, none is available for release.
In 1988, Emilie A. Marcus, Thomas G. Nolen, Catherine H. Rankin, and
Thomas J. Carew published the multiprocess theory to explain disha-
bituation and sensitization in the sea hare, Aplysia. On the basis of their ex-
periments using habituated sea hares that were subjected to different stim-
uli, they concluded that dishabituation and sensitization do not always
occur together; further, they decided, there are three factors to be consid-
ered: dishabituation, sensitization, and inhibition.
Habituation Studies
Habituation studies have utilized a wide variety of approaches, ranging
from the observation of intact organisms carrying out their normal activi-
ties in their natural surroundings to the laboratory observation of individ-
ual nerve cells. With different types of studies, very different aspects of ha-
bituation and sensitization can be investigated. Surveying the animal
kingdom in 1930, G. Humphrey concluded that habituation-like behavior
exists at all levels of life, from the simple one-celled protozoans to the
multicelled, complex mammals.
E. N. Sokolov, a compatriot of Ivan P. Pavlov, used human subjects in the
laboratory. In 1960, he reported on the results of his studies, which in-
volved sensory integration, the makeup of the orientation reflex (which he
322
credited Pavlov with introducing in 1910), a neuronal model and its role in
the orientation reflex, and the way that this neuronal model could be used
to explain the conditioned reflex. Sokolov measured changes in the diame-
ter of blood vessels in the head and finger, changes in electrical waves
within the brain, and changes in electrical conductivity of the skin. By low-
ering the intensity of a tone to which human subjects had been habituated,
Sokolov demonstrated that habituation was not the result of fatigue, be-
cause subjects responded to the lower-intensity tone with the startle or ori-
entation reflex just as they would when a new stimulus was introduced.
Sokolov concluded that the orientation response (which is related to sensi-
tization) and habituation are the result of the functioning of the reticular
network of the brain and central nervous system. Sokolov emphasized that
the orientation response was produced after only the first few exposures to
a particular stimulus, and it increased the discrimination ability of internal
organizers. The orientation response was an alerting command. Heat,
cold, electric shock, and sound were the major stimuli that he used in these
studies.
E. R. Kandell used the sea hare, Aplysia, in his habituation-sensitization
studies. Aplysia is a large sluglike mollusk, with a sheetlike, shell-produc-
ing body covering, the mantle. Aplysia has a relatively simple nervous sys-
tem and an easily visible gill-withdrawal reflex. (The gill is withdrawn into
the mantle shelf.) Early habituation-sensitization experiments dealt with
withdrawal or absence of gill withdrawal. Later experiments measured
electrical changes that occurred within the nerve cells that controlled gill
movement. These were followed by studies which demonstrated that the
gap (synapse) between the receptor nerve cell (sensory neuron) and the
muscle-moving nerve cell (motor neuron) was the site where habituation
and dishabituation occurred and that neurohormones such as acetylcho-
line and serotonin played essential roles in these processes. Kandell called
the synapse the “seat of learning.”
Charles Sherrington used spinal animals in which the connection be-
tween the brain and the spinal nerve cord had been severed. Sherrington
demonstrated that habituation-sensitization could occur within the spinal
nerve cord even without the participation of the brain. Pharmaceuticals
have also been used in habituation-sensitization studies. Michael Davis
and Sandra File used neurotransmitters such as serotonin and norepineph-
rine to study modification of the startle (orientation) response.
Habituation studies conducted in the laboratory enable researchers to
control variables such as genetic makeup, previous experiences, diet, and
the positioning of subject and stimulus; however, they lack many of the
background stimuli present in the field. In her field studies of the chimpan-
323
zees of the Gombe, Jane Goodall used the principles of habituation to de-
crease the distance between herself and the wild champanzees until she
was able to come close enough to touch and be accepted by them. The
field-experimental approach capitalizes on the best of both laboratory
and field techniques. In this approach, a representative group of organisms
that are in their natural state and habitat are subjected to specific, known
stimuli.
Learning to Survive
Habituation is necessary for survival. Many stimuli are constantly imping-
ing upon all living things; since it is biologically impossible to respond si-
multaneously to all of them, those which are important must be dealt with
immediately. It may be a matter of life or death. Those which are unimpor-
tant or irrelevant must be ignored.
Cell physiologists and neurobiologists have studied the chemical and
electrical changes that occur between one nerve cell and another and be-
tween nerve and muscle cells. The results of those studies have been useful
in understanding and controlling these interactions as well as in providing
insights for therapies. Psychologists utilize the fruits of habituation studies
to understand and predict, modify, and control the behavior of intact or-
ganisms. For example, knowing that bulls serving as sperm donors habitu-
ate to one cow or model and stop discharging sperm into it, the animal psy-
chologist can advise the semen collector to use a different cow or model or
simply to move it to another place—even as close as a few yards away.
Conservationists and wildlife protectionists can apply the principles of
habituation to wild animals, which must live in increasingly closer contact
with one another and with humans, so that both animal and human pop-
ulations can survive and thrive. For example, black-backed gulls, when
establishing their nesting sites, are very territorial. Males which enter the
territory of another male gull are rapidly and viciously attacked. After ter-
ritorial boundaries are established, however, the males in contiguous terri-
tories soon exhibit “friendly enemy” behavior: They are tolerant of the
proximity of other males that remain within their territorial boundaries.
This has been observed in other birds as well as in fighting fish.
Walter Lener
See also: Altruism; Communication; Defense mechanisms; Displays;

Ethology; Herbivores; Hierarchies; Insect societies; Isolating mechanisms;
Mammalian social systems; Migration; Mimicry; Omnivores; Pheromones;
Poisonous animals; Predation; Reproductive strategies; Territoriality and
aggression.
324

Alcock, John. Animal Behavior: An Evolutionary Approach. 6th ed. Sunder-
land, Mass.: Sinauer Associates, 1998.
Alkon, Daniel L. “Learning in a Marine Snail.” Scientific American 249 (July,
1983): 70-84.
Barash, David P. Sociobiology and Behavior. New York: Elsevier, 1982.
Drickamer, Lee C., Stephen H. Vessey, and Doug Meikle. Animal Behavior:
Mechanisms, Ecology, Evolution. 4th ed. Dubuque, Iowa: Wm. C. Brown,
1996.
Eckert, Roger, and David Randall. Animal Physiology. 4th ed. San Francisco:
Gould, James L. Ethology: The Mechanisms and Evolution of Behavior. New
York: W. W. Norton, 1980.
Gould, James L., and Peter Marler. “Learning by Instinct.” Scientific Ameri-
can 256 (January, 1987): 74-85.
Halliday, Tim, ed. Animal Behavior. Norman: University of Oklahoma Press,
1994.
Klopfer, P. H., and J. P. Hailman. An Introduction to Animal Behavior: Ethol-
ogy’s First Century. Englewood Cliffs, N.J.: Prentice-Hall, 1967.
Slater, P. J. B., and T. R. Halliday, eds. Behavior and Evolution. New York:
325
HERBIVORES
Types of ecology: Behavioral ecology; Ecoenergetics
Herbivores, animals which eat only plants, include insects and other arthropods,
fish, birds, and mammals. They keep plants from overgrowing and are food for car-
nivores or omnivores.
H erbivores are animals whose diets consist entirely of plants. They oc-
cupy one of the major trophic levels and have two ecological func-
tions. First, they eat plants and keep them from overgrowing. Second, they
are food for carnivores, which subsist almost entirely upon their flesh, and
omnivores, which eat both plants and animals. Herbivores live on land or
in oceans, lakes, and rivers. They can be insects, other arthropods, fish,
birds, or mammals.
Insect Herbivores
Insects are the largest animal class, with approximately one million spe-
cies. Fossils show their emergence 400 million years ago. Insects occur
worldwide, from pole to pole, on land and in fresh or salt water. They are
the best developed invertebrates, except for some mollusks. They mature
by metamorphosis, passing through at least two dissimilar stages before
adulthood. Metamorphosis can take up to twenty years or may be com-
plete a week after an egg is laid.
Many insects are herbivores. Some feed on many different plants, oth-
ers depend on one plant variety or a specific plant portion, such as leaves
or stems. Relationships between insects and the plants they eat are fre-
quently necessary for plant growth and reproduction. Among the insect
herbivores are grasshoppers and social insects such as bees.
Artiodactyls
Artiodactyls are another type of herbivore—hoofed mammals, including
cattle, pigs, goats, giraffes, deer, antelope, and hippopotamuses. Artiodac-
tyls walk on two toes. Their ancestors had five, but evolution removed the
first toe and the second and fifth toes are vestigial. Each support toe—the
third and fourth—ends in a hoof. The hippopotamus, unique among artio-
dactyls, stands on four toes of equal size and width.
What makes artiodactyls herbivorous is the fact that they lack upper in-
cisor and canine teeth, whereas pads in their upper jaws help the lower
teeth grind food. Domesticated artiodactyls include bovids such as cattle,
326
Herbivores
sheep, and goats. Many artiodactyls, such as antelope, cattle, deer, goats,
and giraffes, are ruminants. They chew and swallow vegetation, which en-
ters the stomach for partial digestion, is regurgitated, chewed again, and
reenters the stomach for more digestion. This maximizes nutrient intake
from food. Members of the deer family, which include approximately 40
species from the seven-foot-tall moose to the one-foot-tall pudu, are
hoofed ruminants that inhabit many continents and biomes: Asia, Europe,
the Americas, and North Africa; woods, prairies, swamps, mountains, and
tundra. These animals eat the twigs, leaves, bark, and buds of bushes and
saplings, and grasses.
Most antelope, a group of approximately 150 ruminant species, are Af-
rican, although some are European or Asian. They live on plains, marshes,
deserts, and forests, eating grass, twigs, buds, leaves, and bark. In Asia, Si-
berian saigas and goat antelope (takin) inhabit mountain ranges. Chamois
goat antelope live in Europe’s Alps.There are no true antelope in North
America, where their closest relatives are pronghorns and Rocky Moun-
tain goats (goat-antelope with both goat and antelope anatomic features).
The smallest antelope, the dik-dik, is rabbit-sized. Elands, the largest ante-
lope, are ox-sized.
Giraffes and hippos are artiodactyls that inhabit the dry, tree-scattered
land south of the Sahara in Africa. Giraffes rarely graze and can go for
Deer are herbivores, eating grasses and the tender buds, shoots, bark, and twigs of
trees. The boundary between forest and field offers them the widest range of food
choices. (PhotoDisc)
327
Herbivores
months without drinking, getting most of their water from the leaves they
eat, because it is difficult for them to reach the ground or the surface of a
river with their mouths. By contrast, the short-legged, stocky hippos are
semiaquatic, spending most daylight hours nearly submerged eating
aquatic plants. At night they eat land plants.
Aquatic Herbivores
Fish are aquatic vertebrates, having gills, scales, and fins. They include
rays, lampreys, sharks, lungfish, and bony fish. The earliest vertebrates,
500 million years ago, were fish. They comprise more than 50 percent of all
vertebrates and have several propulsive fins: dorsal fins along the central
back; caudal fins at tail ends; and paired pectoral and pelvic fins on sides
and belly. Fish inhabit lakes, oceans, and rivers, even in Arctic and Antarc-
tic areas. Most marine fish are tropical. The greatest diversity of freshwater
species is found in African and rain forest streams.
Ecological Importance
It is clear that wild herbivores are ecologically important to food chains.
This is because they eat plants, preventing their overgrowth, and they are
eaten by carnivores and omnivores. Domesticated herbivores—cattle,
sheep and goats, used for human sustenance—account for three to four bil-
lion living creatures. Future production of better strains of domesticated
herbivores via recombinant deoxyribonucleic acid (DNA) research may
cut the numbers of such animals killed to meet human needs. Appropriate
species conservation should maintain the present balance of nature and
sustain the number of wild herbivore species living on earth.
Sanford S. Singer
See also: Balance of nature; Food chains and webs; Omnivores; Predation;

Gerlach, Duane, Sally Atwater, and Judith Schnell. Deer. Mechanicsburg,
Pa.: Stackpole Books, 1994.
Gullan, P. J., and P. S. Cranston. Insects: An Outline of Entomology. 2d ed.
Malden, Mass.: Blackwell Science, 2000.
Olsen, Sandra L., ed. Horses Through Time. Boulder, Colo.: Roberts Rine-
hart, 1996.
Rath, Sara. The Complete Cow. Stillwater, Minn.: Voyageur Press, 1990.
Shoshani, Jeheskel, and Frank Knight. Elephants: Majestic Creatures of the
Wild. Rev. ed. New York: Checkmark Books, 2000.
328
HIERARCHIES
Hierarchies, systems of establishing dominance and subordination, are important

in maintaining social order in many species of animals.
A ll animal species strive for their share of fitness. In this struggle for re-
productive success, individuals that make up a population often
compete for essential resources such as food, mates, or nesting sites. In
many species, competition over resources may lead to fighting among in-
dividuals. Fighting, however, can be costly to the individuals involved.
The loser may suffer real injury or even death, and the winner has to ex-
pend energy and still may suffer an injury. In order to prevent constant
fighting over resources, many animal species have adopted a system of
what sociobiologists call a dominance hierarchy or social hierarchy. The
dominance hierarchy is a set of aggression-submission relationships among
the animals of a population. With an established system of dominance, the
subordinate individuals will acquiesce rather than compete with the domi-
nant individuals for resources.
Dominance and Subordinance

To be dominant is to have the priority of access to the essential resources of
life and reproduction. In almost all cases, the superior dominant animals
will displace the subordinates from food, mates, and nest sites. In the mat-
ter of obtaining food, for example, wood pigeons are flock feeders. The
dominant pigeons are always found near the center of the flock when feed-
ing and feed more quickly than the subordinate birds at the edge of the
flock. The birds at the edge of the flock accumulate less food and often ob-
tain just enough to sustain them through the night. Among sheep and rein-
deer, the lowest-ranking females are also the worst-fed animals and among
the poorest of mothers. Baby pigs compete for teat position on the mother
and once established will maintain that position until weaning. Those pig-
lets that gain access to the most anterior teats will weigh more at weaning
than those who have to settle for posterior teat positions. In gaining access
to mates, one study with laboratory mice has shown that while the domi-
nant males constituted only one third of the male population, they sired 92
percent of the offspring.
Life is still not all that hopeless for the subordinates. Oftentimes the
loser in the battle for dominance is given a second chance, and in some of
329
Hierarchies
the more social species, the subordinate only has to await its turn to rise in
the hierarchy. In some species, cooperation among subordinate groups, es-
pecially kin groups, can lead to the formation of a new colony and a new
opportunity to establish dominance. In other species, it may well be ad-
vantageous for the subordinate to stay with the group. For example, indi-
vidual baboons and macaques will not survive very long if they are away
from the group’s sleeping area, and they will have no opportunity to repro-
duce. It has been shown that even a low-ranking male eats well if he is part
of a troop, and he may occasionally have the opportunity to mate. In addi-
tion, the dominant male will eventually lose prowess, and the subordinate
will have a chance to move up in the dominance hierarchy.
Types of Hierarchies
The simplest possible type of hierarchy is a despotism, in which one indi-
vidual rules over all other members of the group and no rank distinctions
are made among the subordinates. Hierarchies more frequently contain
multiple ranks in a more or less linear fashion. An alpha individual domi-
nates all others, a beta individual is subordinate to the alpha but dominates
all others, and so on down to the omega individual at the bottom, who is
dominated by all of the others. Sometimes, the network is complicated by
triangular or other circular relationships in which two or three individuals
might be at the same dominance level. Such relationships appear to be less
stable than despotisms or linear orders.
Nested hierarchies are often observed in some animal species. Societies
that are divided into groups can display dominance both within and be-
tween the various components. For example, white-fronted geese establish
a rank order of several subgroups including parents, mated pairs without
young, and free juveniles. These hierarchies are superimposed over the
hierarchy within each of the subgroups. In wild turkeys, brothers estab-
lish a rank order among their brotherhood, but each brotherhood competes
for dominance with other brotherhoods on the display grounds prior to
mating.
Formation and Maintenance of Hierarchies

Hierarchies are formed during the initial encounters between animals
through repeated threats and fighting, but once the issue of dominance has
been determined, each individual gives way to its superiors with little or
no hostile exchange. Life in the group may eventually become so peaceful
that the existence of ranking is hidden from the observer until some crisis
occurs to force a confrontation. For example, a troop of baboons can go for
hours without engaging in sufficient hostile exchanges to reveal their rank-
330
Hierarchies
ing, but in a moment of crisis such as a quarrel over food the hierarchy will
suddenly be evident. Some species are organized in absolute dominance
hierarchies, in which the rank orders remain constant regardless of the cir-
cumstances. Status within an absolute dominance hierarchy changes only
when individuals move up or down in rank through additional interaction
with their rivals. Other animal societies are arranged in relative dominance
hierarchies. In these arrangements, such as with crowded domestic house
cats, even the highest-ranking individuals acquiesce to subordinates when
the latter approach a point that would normally be too close to their per-
sonal sleeping space.
The stable, peaceful hierarchy is often supported by status signs. In
other words, the mere actions of the dominant individual advertise his
dominance to the other individuals. The leading male in a wolf pack can
control his subordinates without a display of excessive hostility in the
great majority of cases. He advertises his dominance by the way he holds
his head, ears, and tail, and the confident face-forward manner in which he
approaches other members of his pack. In a similar manner, the dominant
rhesus monkey advertises his status by an elaborate posture which in-
cludes elevated head and tail, lowered testicles, and slow, deliberate body
movements accompanied by an unhesitating but measured scrutiny of
other monkeys he encounters. Animals use not only visual signals to ad-
vertise dominance but also acoustic and chemical signals. For example,
dominant European rabbits use a mandibular secretion to mark their terri-
tory.
Uses of Hierarchies
A stable dominance hierarchy presents a potentially effective united front
against strangers. Since a stranger represents a threat to the status of each
individual in the group, he is treated as an outsider. When expelling an in-
truder, cooperation among individuals within the group reaches a maxi-
mum. Chicken producers have long been aware of this phenomenon. If a
new bird is introduced to the flock, it will be subjected to attacks for many
days and be forced down to the lowest status unless it is exceptionally vig-
orous. Most often, it will simply die with very little show of fighting back.
An intruder among a flock of Canada geese will be met with the full range
of threat displays and repeated mass approaches and retreats.
In some primate societies, the dominant animals use their status to stop
fighting among subordinates. This behavior has been observed in rhesus
and pig-tailed macaques and in spider monkeys. This behavior has been
observed even in animal societies, such as squirrel monkeys, that do not
exhibit dominance behavior. Because of the power of the dominant indi-
331
Hierarchies
vidual, relative peace is observed in animal societies organized by despo-

tisms, such as hornets, paper wasps, bumblebees, and crowded territorial
fish and lizards. Fighting increases significantly among the equally ranked
subordinates as they vie for the dominant position when the dominant ani-
mal is removed.
Young males are routinely excluded from the group in a wide range of
aggressively organized mammalian societies such as baboons, langur mon-
keys, macaques, elephant seals, and harem-keeping ungulates. At best,
these young males are tolerated around the fringes of the group, but many
are forced out of the group and either join bachelor herds or wander as soli-
tary nomads. As would be expected, these young males are the most ag-
gressive and troublesome members of the society. They compete with one
another for dominance within their group and often unite into separate
bands that work together to reduce the power of the dominant males.
Males in the two groups show different behaviors. Among the Japanese
macaques, the dominant males stay calm and aloof when introduced to a
new object so as to not risk loss of their status, but the females and young
males will explore new areas and examine new objects.
D. R. Gossett

Ethology; Habituation and sensitization; Herbivores; Insect societies; Iso-
lating mechanisms; Mammalian social systems; Migration; Mimicry; Om-
nivores; Pheromones; Poisonous animals; Predation; Reproductive strate-
gies; Territoriality and aggression.

Barash, David P. Sociobiology and Behavior. 2d ed. New York: Elsevier, 1982.
Campbell, Neil A., Lawrence G. Mitchell, and Jane B. Reece. Biology: Con-
cepts and Connections. 3d ed. San Francisco: Benjamin/Cummings, 2000.
Feldhamer, G. A., L. C. Drickamer, S. H. Vessey, and J. F. Merritt. Mammalogy.
Boston: WCB/McGraw-Hill, 1999.
Ridley, Mark. Animal Behavior: An Introduction to Behavioral Mechanisms, De-
velopment, and Ecology. 2d ed. Boston: Blackwell Scientific Publications,
1995.
Wilson, Edward O. Sociobiology: The Abridged Edition. Cambridge, Mass.:
Belknap Press of Harvard University Press, 1980.
Wittenberger, James F. Animal Social Behavior. Boston: Duxbury Press, 1981.
332
HUMAN POPULATION GROWTH
Since the Industrial Revolution of the nineteenth century, human populations

have experienced a period of explosive growth. Overpopulation now poses a real
threat to plant and animal life, ecosystems, and the long-term sustainability of the
earth’s current ecological balance.
J ust eleven thousand years ago, only about five million humans lived on
the earth. The initial population growth was slow, largely because of the
way humans lived—by hunting and gathering. Such a mobile lifestyle lim-
ited the size of families for practical reasons. When simple means of birth
control, often abstention from sex, failed, a woman would elect abortion or,
more commonly, infanticide to limit her family size. Furthermore, a high
mortality rate among the very young, the old, the ill, and the disabled
acted as a natural barrier to rapid population growth.
Agricultural Revolution
It took more than two million years—from the earliest animal considered
to be human, Homo habilis—or about 100,000 years from the time mod-
ern human beings, Homo sapiens sapiens, migrated out of Africa into the rest
of the world, for the world’s population to reach one billion. The second
billion was added in about one hundred years, the third billion in fifty
years, the fourth in fifteen years, the fifth billion in twelve years. By the
close of the twentieth century, the world’s population was exceeding six
billion.
This explosion had become possible with the development of agricul-
ture. A hunting-gathering lifestyle requires a nomadic existence over a
large range of territory, which makes the establishment of infrastructures,
such as permanent housing and long-range food stores, impractical. Agri-
cultural societies, by contrast, can support more people in a limited area
and, because settlements are permanent, can build infrastructures over
time and therefore minimize efforts directed to basic subsistence, such as
the erecting of shelters. Moreover, when humans became sedentary, some
limits on family size were lifted. With the development of agriculture, chil-
dren became an asset to their families by helping with farming and other
chores.
Starting about eleven thousand years ago (5 million people), humans
began to cultivate such plants as barley, lentils, wheat, and peas in the
333
Human population growth
Middle East—an area that today extends from Lebanon and Syria in the
northwest eastward through Iraq to Iran. In doing so, human beings began
to have a profound ecological impact as well. In cultivating and caring for
these crops, early farmers changed the characteristics of these plants, mak-
ing them higher yielding, more nutritious, and easier to harvest. Agricul-
ture spread and first reached Europe approximately six thousand years
ago. By the beginning of the common era (1 c.e.), human population had
grown to about 130 million, distributed all over the earth.
Agriculture might also have originated independently in Africa in one
or more centers. Many crops were domesticated there, including yams,
okra, coffee, and cotton. In Asia, agriculture based on staples such as rice
and soybeans and many other crops such as citrus, mangos, taro, and ba-
nanas was developed. Agriculture was developed independently in the
New World. It began as early as nine thousand years ago in Mexico and
Peru. Christopher Columbus and his followers found many new crops to
bring back to the Old World, including corn, kidney beans, lima beans, to-
matoes, tobacco, chili peppers, potatoes, sweet potatoes, pumpkins and
squashes, avocados, cacao, and the major cultivated species of cotton.
Ecological Impact
For the last five to six centuries, important staple crops have been culti-
vated throughout the world. Wheat, rice, and corn, which provide 60 per-
cent of the calories people consume, are cultivated wherever they will
grow. Other crops, including spices and herbs, have also been brought
under cultivation. One of the results of the agricultural developments—
particularly pronounced since the Green Revolution of high-yield crops in
the mid-twentieth century—has been a tendency toward “monoculture,”
or the reduction of diversity, in crop plants worldwide.
The growing population has also changed the landscape, distribution,
and diversity of plants dramatically. Clear-cutting and deforestation have
driven many species (both plant and animal) to extinction. Relatively little
has been done to develop agricultural practices suitable for tropical re-
gions. As a result, the tropics are being devastated ecologically, with an es-
timated 20 percent of the world’s species likely to be lost by the mid-
twenty-first century.
Industrial Revolution
By 1650, the world population had reached 500 million. The process of in-
dustrialization had begun, bringing about profound changes in the lives of
humans and their interactions with the natural world. With improved liv-
ing standards, lower death rates, and prolonged life expectancies, human
334
World and Urban Population Growth, 1950-2020

8
7.6
7
6.8
6
Population in billions
6.1
5 5.3
5.0
4 4.5
4.1
3.7
3 3.3
3.0
2.6 2.6
2
1.8
1 1.4
1.1
0.8
0
1950 1960 1970 1980 1990 2000 2010 2020
Total world population Urban population
Sources: Data are from U.S. Bureau of the Census International Data Base and John Clarke,
“Population and the Environment: Complex Interrelationships,” in Population and the
Environment (Oxford, England: Oxford University Press, 1995), edited by Bryan
Cartledge.
Note: The world’s population passed 6 billion in the year 2000.
population grew exponentially. By 1999, there were about 6 billion people,

compared with 2.5 billion in 1950. By 2002, the world population was well
on its way to 7 billion, with an annual growth rate of nearly 100 million.
The processes of industrialization and exponential population growth
combined to multiply the ecological effects of human activity on the bio-
sphere. Human use of fossil fuels, fertilizers, pesticides, global trading
pracitices, agribusiness practices—all these large-scale activities began to
exhibit large-scale effects on both the abiotic and biotic components of eco-
systems everywhere.
Threats to Sustainability
Without effective measures of control, the human population could exceed
the earth’s carrying capacity. Humans are, at present, estimated to con-
sume about 40 percent of the total net products generated via photosynthe-
sis by plants. Human activities have reduced the productivity of earth’s
forests and grasslands by 12 percent. Each year, millions of acres of once-
335
productive land are turned into desert through overgrazing and deforesta-
tion, especially in developing countries. As a result of overfertilization and
aggressive practices in agriculture, topsoil is lost at an annual rate of 24 bil-
lion metric tons. Collectively, these practices caused the destruction of 40
million acres of rain forest each year during the 1960’s and 1970’s and the
extinction of enormous numbers of species.
Through technological innovation and aggressive practices in agricul-
ture, a 2.6-fold increase in world grain production has been achieved since
1950. However, this increase in food output is not nearly enough to feed
the population. Based upon an estimate by the World Bank and the Food
and Agriculture Organization of the United Nations, one out of every five
people is living in absolute poverty, unable to obtain food, shelter, or cloth-
ing dependably. About one out of every ten people receives less than 80
percent of the daily caloric intake recommended by the United Nations. In
countries such as Bangladesh and Haiti and in regions as East Africa, hu-
mans are dying in increasing numbers because of the lack of food. This
food shortage may stem from drought, soil depletion, or soil loss; more of-
ten, famine results from inequitable distribution of resources among popu-
lations. Situations exacerbated by a growing population also pose threats
By the end of the twentieth century, the world’s human population had passed the
6 billion mark, and most of that population lived in or near large urban centers
typified by this street in Barcelona, Spain: with high-rises and other structures de-
signed for high-density living and working accommodations. (PhotoDisc)
336
to the environment, aggravating the problems of acid rains, toxic and haz-
ardous wastes, water shortages, topsoil erosion, ozone layer punctuation,
greenhouse effects, and groundwater contamination.
Ming Y. Zheng
See also: Acid deposition; Biological invasions; Biomagnification; Biopes-

ticides; Deforestation; Erosion and erosion control; Eutrophication; Ge-
netically modified foods; Grazing and overgrazing; Integrated pest man-
agement; Invasive plants; Landscape ecology; Multiple-use approach;
Ocean pollution and oil spills; Ozone depletion and ozone holes; Pesti-
cides; Pollution effects; Population growth; Rangeland; Slash-and-burn
agriculture; Soil; Soil contamination; Urban and suburban wildlife; Waste
management; Wildlife management.

Brown, L. State of the World 2000. New York: W. W. Norton, 2000.
Heiser, C. B., Jr. Seed to Civilization: The Story of Food. 2d ed. San Francisco:
National Research Council. Lost Crops of the Incas: Little-Known Plants of the
Andes with Promise for Worldwide Cultivation. Washington, D.C.: Na-
tional Academy Press, 1990.
Weiner, J. The Next One Hundred Years: Shaping the Future of Our Living
Earth. New York: Bantam Books, 1990.
337
HYDROLOGIC CYCLE
Types of ecology: Ecoenergetics; Ecosystem ecology; Global ecology
The hydrologic cycle, one of the most important geochemical cycles with both
short- and long-range impacts on the biosphere, is a continuous system through
which water circulates through vegetation, in the atmosphere, in the ground, on
land, and in surface water such as rivers and oceans. The sun and earth’s gravity
provide the energy that drives the hydrologic cycle.
T he total amount of water on earth is an estimated 1.36 billion cubic ki-

lometers. Of this water, 97.2 percent is found in the earth’s oceans. The
ice caps and glaciers contain 2.15 percent of the earth’s water. The remain-
der, 0.65 percent, is divided among rivers (0.0001 percent), freshwater and
saline lakes (0.017 percent), groundwater (0.61 percent), soil moisture
(0.005 percent), the atmosphere (0.001 percent), and the biosphere and
groundwater below 4,000 meters (0.0169 percent). While the percentages
of water appear to be small for these water reservoirs, the total volume of
water contained in each is immense.
Evaporation
Evaporation is the process whereby a liquid or solid is changed to a gas.
Heat causes water molecules to become increasingly energized and to
move more rapidly, weakening the chemical force that binds them to-
gether. Eventually, as the temperature increases, water molecules move
from the ocean’s surface into the overlying air. The rate of evaporation is
influenced by radiation, temperature, humidity, and wind velocity.
Each year, about 320,000 cubic kilometers of water evaporate from
oceans. It is estimated that an additional 60,000 cubic kilometers of water
evaporate from rivers, streams, and lakes or are transpired by plants each
year. A total of about 380,000 cubic kilometers of water is evapotranspired
from the earth’s surface every year.
Condensation and Precipitation

Wind may transport the moisture-laden air long distances. The amount of
water vapor the air can hold depends upon the temperature: The higher
the temperature, the more vapor the air can hold. As air is lifted and cooled
at higher altitudes, the vapor in it condenses to form droplets of water.
Condensation is aided by small dust and other particles in the atmosphere.
As droplets collide and coalesce, raindrops begin to form, and precipita-
338
Hydrologic cycle
tion begins. Most precipitation events are the result of three causal factors:
frontal precipitation, or the lifting of an air mass over a moving weather
front; convectional precipitation related to the uneven heating of the earth’s
surface, causing warm air masses to rise and cool; and orographic precipi-
tation, resulting from a moving air mass being forced to move upward
over a mountain range, cooling the air as it rises.
Each year, about 284,000 cubic kilometers of precipitation fall on the
world’s oceans. This water has completed its cycle and is ready to begin a
new cycle. Approximately 96,000 cubic kilometers of precipitation fall
upon the land surface each year. This precipitation follows a number of
different pathways in the hydrologic cycle. It is estimated that 60,000 cubic
kilometers evaporate from the surface of lakes or streams or transpire di-
rectly back into the atmosphere. The remainder, about 36,000 cubic kilome-
ters, is intercepted by human structures or vegetation, infiltrates the soil or
bedrock, or becomes surface runoff.
Interception
In cities, the amount of water intercepted by human structures may ap-
proach 100 percent. However, much urban water is collected in storm sew-
ers or drains that lead to a surface drainage system or is spread over the
land surface to infiltrate the subsoil. Interception loss from vegetation de-
pends upon interception capacity (the ability of the vegetation to collect
and retain falling precipitation), wind speed (the higher the wind speed,
the greater the rate of evaporation), and rainfall duration (the interception
loss will decrease with the duration of rainfall, as the vegetative canopy
will become saturated with water after a period of time). Broad-leaf forests
may intercept 15 to 25 percent of annual precipitation, and a bluegrass lawn
may intercept 15 to 20 percent of precipitation during a growing season.
Transpiration
Plants are continuously extracting soil moisture and passing it into the at-
mosphere through a process called transpiration. Moisture is drawn into
the plant rootlet through osmotic pressure. The water moves through the
plant to the leaves, where it is passed into the atmosphere through the leaf
openings, or stomata. The plant uses less than 1 percent of the soil moisture
in its metabolism; thus, transpiration is responsible for most water vapor
loss from the land in the hydrologic cycle. For example, an oak tree may
transpire 151,200 liters per year.
Overland Flow and Infiltration

When the amount of rainfall is greater than the earth’s ability to absorb it,
339
Hydrologic cycle
The Hydrologic Cycle
Rain clouds
Cloud formation
ing
e fal l
Precipitation wh il
n
Evaporation
ve o ma t i o
n
st r a t i o
tran spi r a t io n
r
f ro m soi l
an
pi
ms
ns
fr o m o c e
t
f
ge
ea
ra
t
ur
S
m
fa
ce
fr o
ru n
Infiltration o ff
Wat er ta bl e
Soil
Zone of Percolation
Rock
saturation Groundwater
Deep percolation Ocean
Source: U.S. Department of Agriculture, Yearbook of Agriculture (Washington, D.C.:

Government Printing Office, 1955).
excess water begins to run off, a process termed overland flow. Overland
flow begins only if the precipitation rate exceeds the infiltration capacity of
the soil. Infiltration occurs when water sinks into the soil surface or into
fractures of rocks; the amount varies according to the characteristics of the
soil or rock and the nature of the vegetative cover. Sandy soils have higher
infiltration rates than clay rock soils. Nonporous rock has an infiltration
rate of zero, and all precipitation that reaches it becomes runoff. The pres-
ence of vegetation impedes surface runoff and increases the potential for
infiltration to occur.
Water infiltrating the soil or bedrock encounters two forces: capillary
force and gravitational force. A capillary force is the tendency of the water
in the subsurface to adhere to the surface of soil or sediment particles. Cap-
illary forces are responsible for the soil moisture a few inches below the
land surface.
The water that continues to move downward under the force of gravity
through the pores, cracks, and fissures of rocks or sediments will eventu-
ally enter a zone of water saturation. This source of underground water is
340
Hydrologic cycle
called an aquifer—a rock or soil layer that is porous and permeable enough
to hold and transport water. The top of this aquifer, or saturated zone, is
the water table. This water is moving slowly toward a point where it is dis-
charged to a lake, spring, or stream. Groundwater that augments the flow
of a stream is called base flow. Base flow enables streams to continue to
flow during droughts and winter months. Groundwater may flow directly
into the oceans along coastlines.
When the infiltration capacity of the earth’s surface is exceeded, over-
land flow begins. Broad, thin sheets of water a few millimeters thick are
called sheet flow. After flowing a few meters, the sheets break up into
threads of current that flow in tiny channels called rills. The rills coalesce
into gullies and, finally, into streams and rivers. Some evaporation losses
occur from the stream surface, but much of the water is returned to the
oceans, thus completing the hydrologic cycle.
Residence Time
Residence time refers to how long a molecule of water will remain in vari-
ous components of the hydrologic cycle. The average length of time that a
water molecule stays in the atmosphere is about one week. Two weeks is
the average residence time for a water molecule in a river, and ten years in
a lake. It would take four thousand years for all the water molecules in the
oceans to be recycled. Groundwater may require anywhere from a few
weeks to thousands of years to move through the cycle. This time period
suggests that every water molecule has been recycled millions of times.
Methods of Study
Several techniques are used to gather data on water in the hydrologic cy-
cle. These data help scientists determine the water budget for different
geographic areas. Together, these data enable scientists to estimate the total
water budget of the earth’s hydrologic cycle.
Scientists have developed a vast array of mathematical equations and
instruments to collect data on the hydrologic cycle. Variations in tempera-
ture, precipitation, evapotranspiration, solar radiation, vegetative cover,
soil and bedrock type, and other factors must be evaluated to understand
the local or global hydrologic cycle.
Precipitation is an extremely variable phenomenon. The United States
has some thirteen thousand precipitation stations equipped with rain
gauges, placed strategically to compensate for wind and splash losses. Pre-
cipitation falling on a given area is determined using a rain-gauge network
of uniform density to determine the arithmetic mean for rainfall in the
area. The amount of water in a snowpack is estimated by snow surveys.
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Hydrologic cycle
The depth and water content of the snowpack are measured and the extent
of the snow cover mapped using satellite photography.
The amount of precipitation lost by interception can be measured and
evaluated. Most often, interception is determined by measuring the
amount above the vegetative canopy and at the earth’s surface. The differ-
ence is what is lost to interception.
The volume of water flowing by a given point at a given time in an open
stream channel is called discharge. Discharge is determined by measuring
the velocity of water in the stream channel, using a current meter. The
cross-sectional area of the stream channel is determined at a specific point
and multiplied by the stream velocity. Automated stream-gauging stations
are located on most streams to supply data for various hydrologic investi-
gations.
The U.S. National Weather Service maintains about five hundred sta-
tions using metal pans, mimicking reservoirs, to measure free-water evap-
oration. Water depths of 17 to 20 centimeters are maintained in the pans.
Errors may result from splashing by raindrops or birds. Because the pans
will heat and cool more rapidly than will a natural reservoir, a pan coeffi-
cient is employed to compensate for this phenomenon. The wind velocity
is also determined. A lake evaporation nomograph determines daily lake
evaporation. The mean daily temperature, wind velocity, solar radiation,
and mean daily dew point are all used in the calculation.
The amount of evapotranspiration can be measured using a lysimeter, a
large container holding soil and living plants. The lysimeter is set outside,
and the initial soil moisture is determined. All precipitation or irrigation is
measured accurately. Changes in the soil moisture storage determine the
amount of evapotranspiration.
Samuel F. Huffman
See also: Balance of nature; Biomass related to energy; Food chains and
webs; Geochemical cycles; Herbivores; Nutrient cycles; Omnivores;
Phytoplankton; Rain forests and the atmosphere; Trophic levels and eco-
logical niches.

Berner, Elizabeth K., and Robert A. Berner. Global Environment: Water, Air,
and Geochemical Cycles. Upper Saddle River, N.J.: Prentice Hall, 1996.
Moore, J. W. Balancing the Needs of Water Use. New York: Springer-Verlag,
1989.
Viessman, Warren, Jr., and Gary L. Lewis. Introduction to Hydrology. 4th ed.
Glenview, Ill.: HarperCollins, 1995.
342
INSECT SOCIETIES
Types of ecology: Behavioral ecology; Population ecology
Ants, termites, and many kinds of bees and wasps live in complex groups known as
insect societies. Studies of such societies have enriched scientific knowledge about
some of the most successful species on earth and have provided insights into the bi-
ological basis of social behavior in other animals.
M any of the most robust, thriving species today owe their success in
great part to benefits that they reap from living in organized groups
or societies. Nowhere are the benefits of group living more clearly illus-
trated than among the social insects. Edward O. Wilson, one of the fore-
most authorities on insect societies, estimates that more than twelve thou-
sand species of social insects exist in the world today. This number is
equivalent to all the species of known birds and mammals combined. Al-
though insect societies have reached their pinnacle in bees, wasps, ants,
and termites, many insects show intermediate degrees of social organiza-
tion—providing insights regarding the probable paths of the evolution of
sociality.
Scientists estimate that eusociality has evolved at least twelve times:
once in the Isoptera, or termites, and eleven separate times in the Hymen-
optera, comprising ants, wasps, and bees. In addition, one group of aphids
has been found to have a sterile soldier caste. Although the eusocial species
represent diverse groups, they all show a high degree of social organi-
zation and possess numerous similarities, particularly with regard to di-
vision of labor, cooperative brood care, and communication among indi-
viduals.
Ants
The organization of a typical ant colony is representative, with minor mod-
ifications, of all insect societies. A newly mated queen, or reproductive fe-
male, will start a new ant colony. Alone, she digs the first nest chambers
and lays the first batch of eggs. These give rise to grublike larvae, which are
unable to care for themselves and must be nourished from the queen’s
own body reserves. When the larvae have reached full size, they undergo
metamorphosis and emerge as the first generation of worker ants. These
workers—all sterile females—take over all the colony maintenance duties,
including foraging outside the nest for food, defending the nest, and clean-
ing and feeding both the new brood and the queen, which subsequently
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Insect societies
becomes essentially an egg-laying machine. For a number of generations,

all eggs develop into workers and the colony grows. Often, several types of
workers can be recognized. Besides the initial small workers, or minor
workers, many ant species produce larger forms known as major workers,
or soldiers. These are often highly modified, with large heads and jaws,
well suited for defending the nest and foraging for large prey. Food may in-
clude small insects, sugary secretions of plants or sap-feeding insects, or
other scavenged foods. After several years, when the colony is large
enough, some of the eggs develop into larger larvae that will mature into
new reproductive forms: queens and males. Males arise from unfertilized
eggs, while new queens are produced in response to changes in larval nu-
trition and environmental factors. These sexual forms swarm out of the
nest in a synchronized fashion to mate and found new colonies of their
own.
Bees and Wasps

With minor modifications, the same pattern occurs in bees and wasps.
Workers of both bees and wasps are also always sterile females, but they
differ from ants in that they normally possess functional wings and lack a
fully differentiated soldier caste. Wasps, like ants, are primarily predators
and scavengers; bees, however, have specialized on pollen and plant nec-
tar as foods, transforming the latter into honey that is fed to both nestmates
and brood. The bias toward females reflects a feature of the biology of the
Hymenoptera that is believed to underlie their tendency to form complex
societies. All ants, wasps, and bees have an unusual form of sex determina-
tion in which fertilized eggs give rise to females and unfertilized eggs de-
velop into males. This type of sex determination, known as haplodiploidy,
generates an asymmetry in the degree of relatedness among nestmates. As
a consequence, sisters are more closely related to their sisters than they are
to their own offspring or their brothers. Scientists believe that this pro-
vided an evolutionary predisposition for workers to give up their own per-
sonal reproduction in order to raise sisters—a form of natural selection
known as kin selection.
Termites
The termites, or Isoptera, differ from the social Hymenoptera in a number of
ways. They derive from a much more primitive group of insects and have
been described as little more than “social cockroaches.” Instead of the
strong female bias characteristic of the ants, bees, and wasps, termites have
regular sex determination; thus, workers have a fifty-fifty sex ratio. Addi-
tionally, termite development lacks complete metamorphosis. Rather, the
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Insect societies
young termites resemble adults in form from their earliest stages. As a con-
sequence of these differences, immature forms can function as workers
from an early age, and—at least among the lower termites—they regularly
do so.
Termites also differ from Hymenoptera in their major mode of feeding.
Instead of feeding on insects or flowers, all termites feed on plant material
rich in cellulose. Cellulose is a structural carbohydrate held together by
chemical bonds that most animals lack enzymes to digest. Termites have
formed intimate evolutionary relationships with specialized microorgan-
isms—predominantly flagellate protozoans and some spirochete bacte-
ria—that have the enzymes necessary to degrade cellulose and release its
food energy. The microorganisms live in the gut of the termite. Because
these symbionts are lost with each molt, immature termites are dependent
upon gaining new ones from their nestmates. They do this by feeding on
fluids excreted or regurgitated by other individuals, a process known as
trophallaxis. This essential exchange of materials also includes, along with
food, certain nonfood substances known as pheromones.
Pheromones, by definition, are chemicals produced by one individual
of a species that affect the behavior or development of other individuals of
the same species that come in contact with them. Pheromones are well doc-
umented throughout the insect world, and they play a key role in commu-
nication between members of nonsocial or subsocial species. Moth mating
attractants provide a well-studied example. Pheromones are nowhere
better developed than among the social insects. They not only appear to in-
fluence caste development in the Hymenoptera and termites but also permit
immediate communication among individuals. Among workers of the fire
ant (Solenopsis saevissima), chemical signals have been implicated in con-
trolling recognition of nestmates, grooming, clustering, digging, feeding,
attraction or formation of aggregations, trail following, and alarm behav-
ior. Nearly a dozen different glands have been identified which produce
some chemical in the Hymenoptera, although the exact function of many of
these chemicals remains unknown.
In addition to chemical communication, social insects may share infor-
mation in at least three other ways: by tactile contact, such as stroking or
grasping; by producing sounds, including buzzing of wings; and by em-
ploying visual cues. Through combinations of these senses, individuals
can communicate complex information to nestmates. Indeed, social insects
epitomize the development of nonhuman language. One such language,
the “dance” language of bees, which was unraveled by Karl von Frisch and
his students, provides one of the best-studied examples of animal behav-
ior. In the waggle dance, a returning forager communicates the location of
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Insect societies
a food resource by dancing on the comb in the midst of its nestmates. It can
accurately indicate the direction of the flower patch by incorporating the
relative angles between the sun, the hive, and the food. Information about
distance, or more precisely the energy expended to reach the food source,
is communicated in the length of the run. Workers following the dance are
able to leave the nest and fly directly to the food source, for distances in ex-
cess of one thousand meters.
Benefits of Cooperation
Living in cooperative groups has provided social insects with opportuni-
ties not available to their solitary counterparts. Not only can more individ-
uals cooperate in performing a given task, but also several quite different
tasks may be carried out simultaneously. The benefits from such coopera-
tion are considerable. For example, group foraging allows social insects to
increase the range of foods they can exploit. By acting as a unit, species
such as army ants can capture large insects and even fledgling birds.
A second benefit of group living is in nest building. Shelter is a primary
need for all animals. Most solitary species use naturally occurring shelters
or, at best, build simple nests. By cooperating and sharing the effort, social
insects are able to build nests that are quite elaborate, containing several
kinds of chambers. Wasps and bees build combs, or rows of special cells,
for rearing brood and storing food. Subterranean termites can construct
mounds more than six meters high, while others build intricate covered
nests in trees. Mound-building ants may cover their nests with a thatch
that resembles, in both form and function, the thatched roofs of old Euro-
pean dwellings. Colonial nesting provides two additional benefits. First, it
enhances defense. By literally putting all of their eggs in one basket, social
insects can centralize and share the guard duties. The effectiveness of this
approach is attested by one’s hesitation to stir up a hornet’s nest.
Nest construction also provides the potential to maintain homeostasis,
the ability to regulate the environment within a desirable range. Virtually
all living creatures maintain homeostasis within their bodies, but very few
animals have evolved the ability to maintain a constant external living en-
vironment. In this respect, insect societies are similar to human societies.
Workers adjust their activities to maintain the living environment within
optimal limits. Bees, for example, can closely regulate the internal temper-
ature of a hive. When temperatures fall below 18 degrees Celsius, they be-
gin to cluster together, forming a warm cover of living bees to protect the
vulnerable brood stages. To cool the hive in hot weather, workers initially
circulate air by beating their wings. If further cooling is needed, they resort
to evaporative cooling by regurgitating water throughout the nest. This
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Insect societies
Social insects such as bees,

wasps, termites, and ants
are known for combining
their efforts to build
protective structures that
allow them to live and
reproduce in relative safety.
This mound, for example,
houses thousands of ants.
(Corbis)
water evaporates with wing fanning and serves to cool the entire hive.
Other social insects rely on different but equally effective methods. Some
ants, and especially termites, build their nests as mounds in the ground,
with different temperatures existing at different depths. The mound nests
of the African termite, Macrotermes natalensis, are an impressive engineer-
ing feat. They are designed to regulate both temperature and air flow
through complex passages and chambers, with the mound itself serving as
a sophisticated cooling tower.
Finally, group living allows the coordination of the efforts of individu-
als to accomplish complex tasks normally restricted to the higher verte-
brates. The similarities between insect societies and human society are
striking. An insect society is often referred to as a superorganism, reflect-
ing the remarkable degree of coordination between individual insects. In-
dividual workers have been likened to cells in a body, and castes to tissues
or organs that perform specialized functions. Insect societies are not im-
mortal; however, they often persist in a single location for periods similar
to the life spans of much larger animals. The social insects have one of the
347
Insect societies
most highly developed symbolic languages outside human cultures. Fur-

ther, social insects have evolved complex and often mutually beneficial in-
teractions with other species to a degree unknown except among human
beings. Bees are inseparably linked with the flowers they feed upon and
pollinate. Ants have actually developed agriculture of a sort with their fun-
gus gardens and herds of tended aphids. On a more sobering note, ants are
the only nonhuman animals that are known to wage war. These striking
similarities with human societies have led researchers to study social in-
sects to learn about the biological basis of social behavior and have led to
the development of a new branch of science known as sociobiology.
Studying Insect Societies

Because of the diversity of questions that investigators have addressed re-
garding insect societies, many methods of scientific inquiry have been em-
ployed. In Karl von Frisch’s experiments, for example, basic behavioral ob-
servations were coupled with simple but elegant experimental design to
unravel the dance language of bees. The bees were raised in an observation
colony. This was essentially a large hive housed between plates of glass so
that an observer could watch the behavior of individual bees. Researchers
followed specific workers by marking them with small numbers placed on
the abdomen or thorax. Sometimes the entire observation hive was placed
within a small, darkened shed to simulate more closely the conditions
within a natural hive.
Bees learned to find an artificial “flower”—a glass dish filled with a
sugar solution. Brightly colored backgrounds and odors such as pepper-
mint oil were added to the sugars to provide specific cues for the bees to as-
sociate with the reward. Feeding stations were set up at fixed distances; ob-
servers could follow the exact movements of known individuals both at
the feeder and at the hive. In this way, von Frisch was able to describe sev-
eral types of dances (the round dance for near food sources, the waggle
dance for feeding stations that were farther from the hive) and show that a
returning bee could share information regarding the location and quality
of a source with her nestmates. Scientists subsequently have developed ro-
bot bees that can be operated by remote control to perform different combi-
nations of dance behaviors. This allows them to determine which parts of
the dance actually convey the coded information.
The investigation of forms of chemical communication requires appli-
cation of a variety of techniques. Chromatography is useful for identifying
the minute amounts of chemical pheromones with which insects commu-
nicate. Chromatography (which literally means “writing with color”) is
particularly suitable for separating mixtures of similar materials. A solu-
348
Insect societies
tion of the mixture is allowed to flow over the surface of a porous solid ma-
terial. Since each component of the mixture will flow at a slightly different
rate, eventually they will become separated or spaced out on the solid ma-
terial. Once the components of the pheromone have been separated and
identified, their activity is assessed separately and in combination using
living insects. Such bioassays allow researchers to determine exactly which
fractions of the chemical generate the highest response.
Other biochemical techniques, such as electrophoresis, have been used
to determine subtle behavioral differences, such as kin discrimination
among hive mates. Each individual carries a complement of enzymes or
proteins that catalyze biological reactions in the body. The structure of such
enzymes is determined by the genetic makeup of the individual, and it var-
ies among individuals. Because enzyme structure is inheritable, however,
much as eye color is, the degree of similarities between the enzymes can be
used as a measure of how closely related two individuals are. The amino
acids composing the enzyme differ in their electrical charges, so different
forms can be separated using the technique of electrophoresis. When a liq-
uid containing their enzymes is subjected to an electrical field, the proteins
with the highest negative charge will move farthest toward the positive
pole. This provides a tool to distinguish close genetic relatives for use in
conjunction with behavioral observations to test, for example, whether
workers can discriminate full sisters from half sisters, or relatives from
nonrelatives, as kin selection theory would predict.
The Success of Social Insects

Social insects are among the most successful groups of animals throughout
the world, especially in the tropics. Although the number of species is low
when compared to all insects (twelve thousand out of more than a million
species), their relative contribution to the community may be unduly large.
In Peru, for example, ants may make up more than 50 percent of the indi-
vidual insects collected at any site.
The study of social insects has provided scientists with new ways of
looking at social behavior in all animals. Charles Darwin described the
evolution of sterile workers in the social insects as the greatest obstacle to
his theory of evolution by natural selection. In attempts to explain this
seeming paradox, William D. Hamilton closely examined the social Hyme-
noptera, where sociality had evolved eleven separate times. Realizing that
the haplodiploid form of sex determination led to sisters being more
closely related to one another than they would be to their own young,
Hamilton developed a far-reaching new theory of social evolution: kin se-
lection, or selection acting on groups of closely related individuals. This
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Insect societies
theory, which provides insights into the evolution of many kinds of seem-
ingly altruistic behaviors, arose primarily from his perceptions regarding
the asymmetrical relatedness of nestmates in the social Hymenoptera. These
insects, then, should be credited with providing the model system that has
led to a subdiscipline of behavioral ecology known as sociobiology, the
study of the biological basis of social behavior. Moreover, given their cen-
tral roles in critical ecological processes such as nutrient cycling and polli-
nation, it would be hard to imagine life without them.
Catherine M. Bristow
See also: Altruism; Animal-plant interactions; Biopesticides; Coevolution;

Communication; Communities: ecosystem interactions; Ethology; Mam-
malian social systems; Mimicry; Pollination; Symbiosis.

Crozier, Ross H., and Pekka Pamilo. Evolution of Social Insect Colonies: Sex
Allocation and Kin Selection. New York: Oxford University Press, 1996.
Frisch, Karl von. The Dance Language and Orientation of Bees. Translated by
L. E. Chadwick. Cambridge, Mass.: Belknap Press, 1967.
Gordon, Deborah M. Ants at Work: How an Insect Society Is Organized. New
York: W. W. Norton, 1999.
Hoelldobler, Bert, and Edward O. Wilson. Journey to the Ants: A Story of Sci-
entific Exploration. Cambridge, Mass.: Belknap Press, 1994.
Ito, Yoshiaki. Behavior and Social Evolution of Wasps: The Communal Aggrega-
tion Hypothesis. New York: Oxford University Press, 1993.
Moffett, Mark W. “Samurai Aphids: Survival Under Siege.” National Geo-
graphic 176 (September, 1989): 406-422.
Prestwich, Glenn D. “The Chemical Defenses of Termites.” Scientific Ameri-
can 249 (August, 1983): 78-87.
Wilson, Edward O. The Insect Societies. Cambridge, Mass.: The Belknap
Press of Harvard University Press, 1971.
_______. Sociobiology: The New Synthesis. Cambridge, Mass.: The Belknap
Press of Harvard University Press, 1975.
_______. Success and Dominance in Ecosystems: The Case of the Social Insects.
Oldendorf/Luhe, Federal Republic of Germany: Ecology Institute,
1990.
350
INTEGRATED PEST MANAGEMENT
Types of ecology: Agricultural ecology; Ecotoxicology; Restoration and
conservation ecology
Integrated pest management (IPM) is the practice of integrating insect, animal, or

plant management tactics, such as chemical control, cultural control, biological
control, and plant resistance, to maintain pest populations below damaging levels
in the most economical and environmentally responsible manner.
I n the past, pest management strategies in agriculture focused primarily

on eliminating all of a particular pest organism from a given field or
area. These strategies depended on the use of chemical pesticides to kill all
the pest organisms. Prior to the twentieth century, farmers used naturally
occurring compounds such as kerosine or pyrethrum for this purpose.
During the second half of the twentieth century, synthetic pesticides began
playing a prominent role in controlling crop pests.
Chemical Effects
After 1939 the use of pesticides such as dichloro-diphenyl-trichloroethane
(DDT) was so successful in controlling pest populations that farmers be-
gan to substitute a heavy dependence on pesticides for sound pest man-
agement strategies. Soon pests in high-value crops became resistant to one
pesticide after another. In addition, outbreaks of secondary pests occurred
because either they developed resistance to the pesticides or the pesticides
killed their natural enemies. Among birds ingesting DDT in the food chain,
eggshells were so thin as to render many eggs unviable, reducing the bird
population. Such impacts of DDT supplied the impetus for chemical com-
panies to develop new pesticides, to which the pests also eventually devel-
oped resistance.
Rationale for IPM

Certain pests have developed resistance to all federally registered materi-
als designed to control them. In addition, many pesticides are toxic to hu-
mans, wildlife, and other nontarget organisms and therefore contribute to
environmental pollution. For these reasons, and because it is very expen-
sive for chemical companies to put a new pesticide on the market, many
producers began looking at alternative strategies such as IPM for manag-
ing pests. The driving forces behind the development of IPM programs are
concern about the contamination of groundwater and other nontarget
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Integrated pest management
sites, adverse effects on nontarget organisms, and development of pesti-

cide resistance. Pesticides will probably continue to play a vital role in pest
management, even in IPM, but it is believed that the role will be greatly di-
minished over time.
An agricultural ecosystem consists of the crop environment and its sur-
rounding habitat. The interactions among soil, water, weather, plants, and
animals in this ecosystem are rarely constant enough to provide the eco-
logical stability of nonagricultural ecosystems. Nevertheless, it is possible
to use IPM to manage most pests in an economically efficient and environ-
mentally friendly manner. IPM programs have been successfully imple-
mented in the cropping of cotton and potatoes, and they are being devel-
oped for other crops.
Developing IPM Programs

There are generally three stages of development associated with IPM pro-
grams, and the speed at which a program progresses through these stages
is dependent on the existing knowledge of the agricultural ecosystem and
the level of sophistication desired. The first phase is referred to as the pesti-
cide management phase. The implementation of this phase requires that the
farmer know the relationship between pest densities and the resulting dam-
age to crops so that the pesticide is not applied excessively. In other words,
farmers do not have to kill all the pests all the time. They must use pesticides
only when the economic damage caused by a number of pest organisms
present on a given crop exceeds the cost of using a pesticide. This practice
alone can reduce the number of chemical applications by as much as half.
The second phase is called the cultural management phase. Implemen-
tation of this phase requires knowledge of the pest’s biology and its rela-
tionship to the cropping system. Cultural management includes such prac-
tices as delaying planting times, rotating crops, altering harvest dates, and
planting resistant cultivars. It is necessary to understand pest responses to
other species as well as abiotic factors, such as temperature and humidity,
in the environment. If farmers know the factors that control population
growth of a particular pest, they may be able to reduce the impact of that
pest on a crop. For example, if a particular pest requires short days to com-
plete development, farmers might be able to harvest the crop before the
pest has a chance to develop.
The third phase is the biological control phase, which involves the use
of biological organisms rather than chemicals to control pests. This is the
most difficult phase to implement because farmers must understand not
only the pest’s biology but also the biology of the pest’s natural enemies
and the degree of effectiveness with which these agents control the pest.
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Integrated pest management
In general, it is not possible to rely completely on biological control

methods. A major requirement in using biological agents is to have suffi-
cient numbers of the control agent present at the same time that the pest
population is at its peak. It is sometimes possible to change the planting
dates so that the populations of the pests and the biological control agents
are synchronized. Also, there is often more than one pest species present at
the same time within the same crop, and it is extremely difficult to control
simultaneously two pests with biological agents.
D. R. Gossett
See also: Biomagnification; Biopesticides; Genetically modified foods; Pesti-

cides; Pollution effects; Soil contamination.

Pedigo, Larry P. Entomology and Pest Management. 4th ed. Upper Saddle
River, N.J.: Prentice Hall, 2002.
Romoser, William S., and John G. Stoffolano. The Science of Entomology. 4th
ed. Boston: McGraw-Hill, 1998.
353
INVASIVE PLANTS
Types of ecology: Ecosystem ecology; Ecotoxicology
Nonnative (also termed “exotic”) fungi and plants that can outcompete native
species are called invasive plants. Invasive plants cause irreversible changes to eco-
systems, threaten plant and animal species, and cost billions of dollars to control.
B etween the damage they cause and the cost of control efforts, invasive
plants cost the United States more than $140 billion every year. For ex-
ample, nearly half of the threatened and endangered plant species listed
for the United States in 1999, 400 of 958, were in peril because of competi-
tion from invasive species. Thus, invasive plants are capable of causing ir-
reparable changes in ecosystems.
Most invasive species in the United States originated in Asia or Europe.
The seeds or spores of these plants are accidentally transported into new
habitats by humans, leaving the plants’ natural enemies and competitors
behind. Without natural biological controls, the alien species can thrive
and outcompete the native flora, driving the native plants toward extinc-
tion and creating a near monoculture of the invader.
Invasive plants are weedy species that grow rapidly, produce large
numbers of long-lived seeds, and frequently have perennial roots, or rhi-
zomes, that enhance asexual propagation. Invasive plants have a variety of
effects on invaded ecosystems. Many invasive species deplete soil mois-
ture and nutrient levels, either by growing more vigorously than native
plants early in the growing season or by being more tolerant of reduced
levels of water and nutrients than are natives. Some invasive species pro-
duce toxic chemicals (allelopathy) that are released into the soil and inhibit
the growth of competitors. By outcompeting native plants, the invader de-
creases species diversity as it replaces many native species. As a result, ani-
mal species dependent on native flora are also affected. Fungi and seed
plants are among the most disruptive invasive plants in the United States
today.
Control Methods
Invasive species are carried to new habitats, either in or on machinery or
organisms, and are usually transported by humans, so prevention is the
most cost-effective method of control. Once an invasive species has en-
tered an area, plant quarantine is an effective first line of defense. For ex-
ample, living plants and animals brought into the United States must pass
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Invasive plants
inspection by the U.S. Department of Agriculture Animal and Plant Health

Inspection Service (APHIS) to ensure that they are not carrying potentially
invasive species. Particular care is taken to ensure that imports from
known areas of infestation are clean of seeds, spores, or propagules.
The next most effective strategy is detection and control of small infesta-
tions. When there is a known threat of invasion, the affected area should be
surveyed periodically and individual plants removed by hand or, in ex-
treme cases, by “spot-spraying” herbicide. Eradication is possible when
the infestation is small.
Once an invasive species becomes established, the only means of man-
agement are expensive chemical or biological controls which, at best, will
only minimize damage. A variety of chemicals may be used to kill invasive
plants. Most chemicals, however, affect a broad spectrum of plants, includ-
ing native species. Biological controls, including natural enemies from the
invasive plant’s native ecosystem, can be more specific but may also be ca-
pable of displacing native species and becoming “invaders.”
Fungi
Many of the most serious plant pathogens are invasive species introduced
into the Americas since the beginning of European settlement. Two classic
examples are Dutch elm disease, caused by the fungi Ophiostoma ulmi and
Ophiostoma novo-ulmi and chestnut blight, caused by the fungus Crypho-
nectria parasitica. At the beginning of the twentieth century, the most com-
mon street tree growing in the cities of the eastern United States was the
Image Not Available
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Invasive plants
American elm. About 1910, the European bark beetle was introduced into
the United States. It was not until the 1930’s that Dutch elm disease was ob-
served in Ohio and a few eastern states. The fungal spores are carried by
the beetles, which burrow under the elm bark. The native elm has little re-
sistance to this fungus, whose spores rapidly germinate and form exten-
sive mycelia within the phloem of the host tree, killing it within a few
years. After its initial contact, the fungus spread throughout the cities and
forests of the East and gradually westward, so that by 1990 nearly all the
native American elm trees in the United States had been killed.
American chestnut was also one of the early dominant trees of the east-
ern U.S. forest. In addition to providing edible fruit, the chestnut became a
commercially important timber tree. Chestnut blight fungus was first re-
ported in 1904 on chestnut trees in the New York Zoological Garden and
quickly began to spread. This infestation led directly to passage of the
Plant Quarantine Act of 1912, the forerunner of APHIS. By 1950 most na-
tive chestnut trees were reduced to minor understory shrubs. Biological
control using virus strains first isolated in Italy show promise for control-
ling the blight.
Terrestrial Green Plants

Virtually all the plants commonly called weeds are foreign invaders that
are difficult, if not impossible, to control. Some of the most severe include
Canada thistle (Circium arvense), leafy spurge (Euphorbia esula), and purple
loosestrife (Lythrum salicaria).
Canada thistle is the most widespread and difficult species of thistle to
control. It was introduced to Canada from Europe in the 1600’s and in 1795
was listed as a noxious weed in Vermont. It is now found in most of the
United States as well as in Canada. Single herbicide applications do not
provide long-term control, and there are no effective biological controls
that do not also attack native species.
Leafy spurge was first reported in Newbury, Massachusetts, in 1827,
where it arrived in ship ballast. By 1900 it had reached the West Coast, and
it now thrives in more than half the states and in Canada. Thirteen species
of insects are approved for biological control, and several herbicides can be
used to control infestations effectively. Sheep and goats will browse on
spurge.
Purple loosestrife was introduced into the United States as an ornamen-
tal plant in the early 1800’s and became established in New England by
1830. Its early spread into the Great Lakes region was by barge and other
canal traffic. Rapid expansion of the pest, particularly in the West, occurred
after 1940, primarily due to the plant’s “escape” from ornamental cultiva-
356
Invasive plants
tion into irrigation projects. It is now found in all the lower forty-eight
states except Florida. At present, there are no effective controls.
Aquatic Green Plants

Invasive plants are not limited to the terrestrial habitat or to vascular
plants. One dramatic example is the alga Caulerpa taxifolia, the so-called
killer alga. This attractive tropical alga was found to be easy to grow in salt-
water aquaria and useful as a secondary food source for herbivorous tropi-
cal fish. It began to be used this way at the Oceanographic Museum of
Monaco in 1982. Two years later, a meter-square patch was found growing
in the Mediterranean Sea, visible from a window of the museum. By 1990
the alga had reached France, and by 1995 it could be found from Spain to
Croatia. Caulerpa produces a number of toxins that inhibit foraging by na-
tive fish, and it is a prolific vegetative reproducer. Fragments of the alga,
stuck on an anchor for example, can start a new infestation wherever the
anchor is next dropped. This species has been discovered in Southern Cali-
fornia, and a related species has become dominant in Sydney Harbor, Aus-
tralia. Other aquatic invasive plants in the United States include the mos-
quito fern (Azolla), the Eurasian water milfoil (Myriophyllum), and the
water hyacinth (Eichhornia crassipes).
See also: Biological invasions; Biomagnification; Eutrophication; Genet-

ically modified foods; Pesticides; Phytoplankton; Pollution effects; Waste
management.

Meinesz, Alexandre. Killer Algae. Chicago: University of Chicago Press,
1999.
Randall, John M., and Janet Marinelli, eds. Invasive Plants: Weeds of the
Global Garden. Brooklyn, N.Y.: Brooklyn Botanic Garden, 1996.
Sheley, Roger L., and Janet K. Petroff. Biology and Management of Noxious
Rangeland Weeds. Corvalis, Oreg.: Oregon State University Press, 1999.
357
ISOLATING MECHANISMS
Types of ecology: Behavioral ecology; Ecosystem ecology; Evolutionary
ecology; Speciation
Isolating mechanisms act to prevent interbreeding and the exchange of genes be-
tween species. The establishment of isolating mechanisms between populations is a
critical step in the formation of new species and ensuring biodiversity.
I solating mechanisms (reproductive isolating mechanisms) prevent in-

terbreeding between species. The term, which was first used by Theo-
dosius Dobzhansky in 1937 in his landmark book Genetics and the Origin of
Species, refers to mechanisms that are genetically influenced and intrinsic.
Geographic isolation can prevent interbreeding between populations, but
it is an extrinsic factor and therefore does not qualify as an isolating mecha-
nism. Isolating mechanisms function only between sexually reproducing
species. They have no applicability to forms that reproduce only by asex-
ual means, as by mitotic fission, stoloniferous or vegetative reproduction,
or egg development without fertilization (parthenogenesis in animals).
Obligatory self-fertilization in hermaphrodites (mainly a phenomenon in
plants, rare in animals) is a distortion of the sexual process that produces
essentially the same results as asexual reproduction. Many lower animals
and protists regularly employ both asexual and sexual means of reproduc-
tion, and the significance of isolating mechanisms in such forms is essen-
tially the same as in normal sexual species.
Premating Mechanisms
Reproductive isolating mechanisms are usually classified into two main
groups. Premating (prezygotic) mechanisms operate prior to mating, or
the release of gametes, and, therefore, do not result in a wastage of the re-
productive potential of the individual. Postmating (postzygotic) mecha-
nisms come into play after mating, or the release of gametes, and could re-
sult in a loss of the genetic contribution of the individual to the next
generation. This distinction is also important in the theoretical sense in that
natural selection should favor genes that promote premating isolation;
those that do not presumably would be lost more often through mis-
matings (assuming that hybrids are not produced, or are sterile or inferior),
and this could lead to a reinforcement of premating isolation.
Ethological (behavioral) isolation is the most important category of
premating isolation in animals. The selection of a mate and the mating pro-
358
Isolating mechanisms
cess depends upon the response of both partners to various sensory cues,
any one of which may be species-specific. Although one kind of sensory
stimulus may be emphasized, different cues may come into play at differ-
ent stages of the pairing process. Visual signals provided by color, pattern,
or method of display are often of particular importance in diurnal animals
such as birds, many lizards, certain spiders, and fish. Sounds, as in male
mating calls, are often important in nocturnal breeders such as crickets or
frogs but are also important in birds. Mate discrimination based on chemi-
cal signals or odors (pheromones) is of fundamental importance in many
different kinds of animals, especially those where visual cues or sound are
not emphasized; chemical cues also are often important in aquatic animals
with external fertilization. Tactile stimuli (touch) often play an important
role in courtship once contact is established between the sexes. Even elec-
trical signals appear to be utilized in some electrogenic fish.
Ecological (habitat) isolation often plays an important role. Different
forms may be adapted to different habitats in the same general area and
may meet only infrequently at the time of reproduction. One species of
deer mouse, for example, may frequent woods, while another is found in
old fields; one fish species spawns in riffles, while another spawns in still
pools. This type of isolation, although frequent and widespread, is often
incomplete as the different forms may come together in transitional habi-
tats. The importance of ecological isolation, however, is attested by the fact
that instances in which hybrid swarms are produced between forms that
normally remain distinct have often been found to be the result of disrup-
tion of the environment, usually by humans. Mechanical isolation is a less-
important type of premating isolation, but it can function in some combi-
nations. Two related animal species, for example, may be mismatched be-
cause of differences in size, proportions, or structure of genitalia.
Finally, temporal differences often contribute to premating isolation.
The commonest type of temporal isolation is seasonal isolation: Species
may reproduce at different times of the year. A species of toad in the east-
ern United States, for example, breeds in the early spring, while a related
species breeds in the late spring, with only a short period of overlap. Dif-
ferences can also involve the time of day, whereby one species may mate at
night and another during the day. Such differences, as in the case of ecolog-
ical isolation, are often incomplete but may be an important component of
premating isolation.
Postmating Mechanisms
If premating mechanisms fail, postmating mechanisms can come into play.
If gametes are released, there still may be a failure of fertilization (inter-
359
sterility). Spermatozoa may fail to penetrate the egg, or even with penetra-
tion there may be no fusion of the egg and sperm nucleus. Fertilization fail-
ure is almost universal between remotely related species (as from different
families or above) and occasionally occurs even between closely related
forms.
If fertilization does take place, other postmating mechanisms may oper-
ate. The hybrid may be inviable (F1 or zygotic inviability). Embryonic de-
velopment may be abnormal, and the embryo may die at some stage, or the
offspring may be defective. In other cases, development may be essen-
tially normal, but the hybrid may be ill-adapted to survive in any available
habitat or cannot compete for a mate (hybrid adaptive inferiority). Even
if hybrids are produced, they may be partially to totally sterile (hybrid ste-
rility). Hybrids between closely related forms are more likely to be fer-
tile than those between more distantly related species, but the correlation is
an inexact one. The causes for hybrid sterility are complex and can involve
genetic factors, differences in gene arrangements on the chromosomes that
disrupt normal chromosomal pairing and segregation at meiosis, and in-
compatibilities between cytoplasmic factors and the chromosomes. If the
hybrids are fertile and interbreed or backcross to one of the parental
forms, a more subtle phenomenon known as hybrid breakdown some-
times occurs. It takes the form of reduced fertility or reduced viability in
the offspring. The basis for hybrid breakdown is poorly understood but
may result from an imbalance of gene complexes contributed by the two
species.
It should be emphasized that in most cases of reproductive isolation
that have been carefully studied, more than one kind of isolating mecha-
nism has been found to be present. Even though one type is clearly of para-
mount importance, it is usually supplemented by others, and should it fail,
others may come into play. In this sense, reproductive isolation can be
viewed as a fail-safe system. A striking difference in the overall pattern of
reproductive isolation between animals and plants, however, is the much
greater importance of premating isolation in animals and the emphasis on
postmating mechanisms in plants. Ethological isolation, taken together
with other premating mechanisms, is highly effective in animals, and
postmating factors usually function only as a last resort.
Field Studies and Experimental Studies

Field studies have often been employed in the investigation of some types
of premating isolating mechanisms. Differences in such things as breeding
times, factors associated with onset of breeding activity, and differences in
habitat distribution or selection of a breeding site are all subject to direct
360
field observation. Comparative studies of courtship behavior in the field or

laboratory often provide clues as to the types of sensory signals that may
be important in the separation of related species.
Mating discrimination experiments carried on in the laboratory have
often been employed to provide more precise information on the role
played by different odors, colors, or patterns, courtship rituals, or sounds
in mate selection. Certain pheromones, for example, which act as sexual at-
tractants, have been shown to be highly species-specific in some insects.
The presence or absence of certain colors or their presentation has been
shown experimentally to be important in mate discrimination in ver-
tebrates as diverse as fish, lizards, and birds. Call discrimination ex-
periments, in which a receptive female is given a choice between recorded
calls of males of her own and another species, have demonstrated the
critical importance of mating call differences in reproductive isolation in
frogs and toads. Synthetically generated calls have sometimes been used to
pinpoint the precise call component responsible for the difference in re-
sponse.
Studies on postmating isolating mechanisms have most often involved
laboratory crosses in which the degree of intersterility, hybrid sterility, or
hybrid inviability can be analyzed under controlled conditions. In in-
stances in which artificial crosses are not feasible, natural hybrids some-
times occur and can be tested. The identification of natural backcross prod-
ucts can attest incomplete postmating, as well as premating isolation.
Instances of extensive natural hybridization are of special interest and
have often been subjected to particularly close scrutiny. Such cases often
throw light on factors that can lead to a breakdown of reproductive isola-
tion. Also, as natural hybridization more often occurs between marginally
differentiated forms in earlier stages of speciation, new insights into the
process of species formation can sometimes be obtained. Finally, such
studies may yield information on the evolutionary role of hybridization,
including introgressive hybridization, the leakage of genes from one spe-
cies into another. Morphological analysis has long been used in such cases,
and chromosomal studies are sometimes appropriate. In recent years,
allozyme analysis by gel electrophoresis has become a routine tool in esti-
mates of gene exchange, and molecular analysis of nuclear deoxyribonu-
cleic acid (DNA), mitochondrial DNA, have been useful. As mitochondria
are normally passed on only maternally, their DNA can also be used to
identify cases in which females of only one of the two species has been in-
volved in the breakdown of reproductive isolation.
Investigations of the role of natural selection in the development and re-
inforcement of reproductive isolation have employed two different ap-
361
proaches. One has involved the measurement of geographic variation in

the degree of difference in some signal character (call, color, or pattern, for
example) thought to function in premating isolation between two species
that have overlapping ranges. If the difference is consistently greater
within the zone of overlap (reproductive character displacement), an argu-
ment can be made for the operation of reinforcement. Another approach
has involved laboratory simulations, usually with the fruit fly Drosophila,
in which some type of selective pressure is exerted against offspring pro-
duced by crosses between different stocks, and measurement is made of
the frequency of mismatings through successive generations. The results
of such studies to this time are contradictory, and the role of selection with
regard to development of reproductive isolation requires further study.
Enhancing Reproductive Efficiency

The efficiency of reproduction in most animals is enhanced immeasurably
by premating isolating mechanisms. Clearly, in animals a random testing
of potential mates without regard to type is totally unacceptable for most
species in terms of reproductive capacity and time and energy resources.
Premating isolation in this sense is a major factor in promoting species di-
versity in animal communities.
Both premating and postmating isolating mechanisms are also critical
to the maintenance of species diversity in that they act to protect the ge-
netic integrity of each form: A species cannot maintain its identity without
barriers that prevent the free exchange of genes with other species. Fur-
thermore, a species functions as the primary unit of adaptation. Every spe-
cies in a community has its own unique combination of adaptive features
that enable it to exploit the resources of its environment and to coexist with
other species with a minimum of competition. The diversity of different
species that can coexist in the same area depends upon the unique “niche”
that each occupies; adaptive features that determine that niche are based
on the unique genetic constitution of each species, and this genetic consti-
tution is protected through reproductive isolation.
The development of reproductive isolating mechanisms is also critical
to the formation of new species (speciation), and ultimately to the develop-
ment of new organic diversity. The most widely accepted, objective, and
theoretically operational concept for a sexual species is the biological spe-
cies concept. Such a species can be defined as population or group of pop-
ulations, members of which are potentially capable of interbreeding but
which are reproductively isolated from other species. The origin of new
species, therefore, depends upon the development of reproductive iso-
lating mechanisms between populations. A major focus of research in
362
evolutionary biology and systematics has been, and continues to be, on the
various factors that influence the development of reproductive isolating
mechanisms.
John S. Mecham
See also: Adaptive radiation; Biodiversity; Clines, hybrid zones, and

introgression; Communication; Convergence and divergence; Evolution:
definition and theories; Gene flow; Mammalian social systems; Natural se-
lection; Nonrandom mating, genetic drift, and mutation; Punctuated equi-
librium vs. gradualism; Reproductive strategies; Speciation.

Dobzhansky, Theodosius. Genetics of the Evolutionary Process. New York:
Columbia University Press, 1970.
Dobzhansky, Theodosius, Francisco J. Ayala, G. Ledyard Stebbins, and
James W. Valentine. Evolution. San Francisco: W. H. Freeman, 1977.
Mayr, Ernst. Populations, Species, and Evolution. Cambridge, Mass.: Harvard
363
LAKES AND LIMNOLOGY
Types of ecology: Aquatic and marine ecology; Biomes; Ecosystem
ecology
Lakes are inland bodies of water that fill depressions in the earth’s surface. They are
generally too deep to allow vegetation to cover the entire surface and may be fresh
or saline. The study of the physical, chemical, climatological, biological, and eco-
logical aspects of lakes is known as limnology.
Geological Origin of Lakes

Several geologic mechanisms can create the closed basins that are needed
to impound water and produce lakes. The most important of these mecha-
nisms include glaciers, landslides, volcanoes, rivers, subsidence, and tec-
tonic processes.
Continental glaciers formed thousands of lakes by the damming of
stream valleys with moraine materials. Glaciers also scoured depressions
in softer bedrock, and these later filled with water to form lakes. Depres-
sions called kettles formed when buried ice blocks melted. Mountain gla-
ciers also produce numerous small, high alpine lakes by plucking away
bedrock. The bowl-shaped depressions that occur as a result of this pluck-
ing are called cirques; lakes that occupy cirques are called tarns. Sometimes
a mountain glacier moves down a valley and carves a series of depressions
along the valley that, from above, look like a row of beads along a string.
When these depressions later fill with water, the lakes are called paternos-
ter lakes, the name coming from their similarity to beads on a rosary.
Landslides sometimes form natural dams across stream valleys. Large
lakes then pond up behind the dam. Volcanoes may produce lava flows
that dam stream valleys and produce lakes. A volcanic explosion crater
may fill with water and make a lake. After an eruption, the area around the
eruption vent may collapse to form a depression called a caldera. Some cal-
deras, such as Crater Lake in Oregon, fill with water. Rivers produce lakes
along their valleys when a tight loop of a meandering channel finally is
eroded through and leaves behind an oxbow lake, isolated from the main
channel. Sediment may accumulate at the mouth of a stream, and the re-
sulting delta may build, bridging across irregularities in the shoreline to
create a brackish coastal lake.
Natural subsidence creates closed basins in areas underlain by soluble
limestones or evaporite deposits. As the underlying limestone is dissolved
away, the earth above collapses to form a cavity (sinkhole), which later fills
364
Lakes and limnology
with water. Finally, large-scale (tectonic) downwarping of the earth’s crust

produces some very large lakes. Large basins form when the crust warps
or sinks downward in response to deep forces. The subsidence produces
very large closed basins that can hold water. A few immense lakes owe
their origins to tectonic downwarping.
Sedimentation
With few exceptions, most lakes exist in relatively small depressions and
serve as the catch basins for sediment from the entire watershed around
them. The natural process of sedimentation ensures that most lakes fill with
sediment before very long periods of geologic time have passed. Lakes with
areas of only a few square kilometers or less will fill within a few tens of
thousands of years. Very large lakes, the inland seas, may endure for more
than ten million years. Human-made lakes and reservoirs have unusually
high sediment-fill rates in comparison with most natural lakes. Human-
made lakes fill with sediment within a few decades to a few centuries.
Lake sediments come from four sources: allogenic clastic materials that
are washed in from the surrounding watershed; endogenic chemical pre-
cipitates that are produced from dissolved substances in the lake waters;
Lakes offer rich habitats for complex ecosystems comprising organisms that live on
land, in water, or in both environments. Depending on their elevation, lakes can
support wildlife typical of taiga, such as freshwater fish and bears, or lower eleva-
tions, such as migratory seabirds. (Digital Stock)
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Lakes and limnology
endogenic biogenic organic materials produced by plants and animals liv-

ing in the lake; and airborne substances, such as dust and pollen, trans-
ported to the lake in the atmosphere.
Allogenic clastic materials are mostly minerals; they are produced
when rocks and soils in the drainage basin are weathered by mechanical
and chemical processes to yield small particles. These particles are moved
downslope by gravity and running water to enter streams, which then
transport them to the lake. Clastic materials also enter the lake via waves,
which erode the materials from the shoreline, and via landslides that di-
rectly enter the lake. In winter, ice formed on the lake can expand and push
its way a few centimeters to 1 meter or so onto the shore. There, the ice may
pick up large particles, such as gravel and cobbles. When spring thaw
comes, waves can remove that ice, together with its enclosed particles, and
float it out onto the lake. The process by which the large particles are trans-
ported out on the lake is called ice-rafting. As the ice melts, the large clastic
particles drop to the bottom; they are termed dropstones when found in
lake sediments. A landslide into a lake or a flood on a stream that feeds into
the lake can produce water heavily laden with sediment. The sediment-
laden water is more dense than clean water and therefore can rush down
and across the lake bottom at speeds sufficient to carry even coarse sand
far out into the lake. These types of deposits are called turbidite deposits.
Endogenic chemical precipitates in freshwater lakes commonly consist
of carbonate minerals (calcite, aragonite, or dolomite) and mineraloids that
consist of oxides and hydroxides of iron, manganese, and aluminum. In
some saline and brine lakes, the main sediments may be carbonates, to-
gether with sulfates such as gypsum (hydrated calcium sulfate), thenardite
(sodium sulfate), or epsomite (hydrated magnesium sulfate), or with chlo-
rides such as halite (sodium chloride) or more complex salts. Of the
endogenic precipitates, calcite is the most abundant. Its precipitation rep-
resents a balance between the composition of the atmosphere and that of
the lake water.
Diatoms are distinctive microscopic algae that produce a frustule (a
kind of shell) made of silica glass that is highly resistant to weathering.
When seen under a high-powered microscope, diatom frustules appear to
be artwork—beautiful and highly ornate saucer- and pen-shaped works of
glass. A tiny spot of lake sediment may contain millions.
A lake’s sediment may contain from less than 1 percent to more than 90
percent organic materials, depending upon the type of lake. Most organic
matter in lake sediments is produced within the lake by plankton and con-
sists of compounds such as carbohydrates, proteins, oils, and waxes that
are made up of organic carbon, hydrogen, nitrogen, and oxygen, with a lit-
366
Lakes and limnology
tle phosphorus. Plankton, with an approximate bulk composition of 36

percent carbon, 7 percent hydrogen, 50 percent oxygen, 6 percent nitro-
gen, and 1 percent phosphorus (by weight), includes microscopic plants
(phytoplankton) and microscopic animals (zooplankton) that live in the
water column. Lakes that are very high in nutrients (eutrophic lakes) com-
monly have heavy blooms of algae, which contribute much organic matter
to the bottom sediment. Terrestrial (land-derived) organic material such as
leaves, bark, and twigs form a minor part of the organic matter found in
most lakes. Terrestrial organic material is higher in carbon and lower in hy-
drogen, nitrogen, and phosphorus than is planktonic organic matter.
Airborne substances usually constitute only a tiny fraction of lake sedi-
ment. The most important material is pollen and spores. Pollen usually
constitutes less than 1 percent of the total sediments, but that tiny amount
is a very useful component for learning about the recent climates of the
earth. Pollen is among the most durable of all natural materials. It survives
attack by air, water, and even strong acids and bases. Therefore, it remains
in the sediment through geologic time. As pollen accumulates in the bot-
tom sediment, the lake serves as a kind of recorder for the vegetation that
exists around it at a given time. By taking a long core of the bottom sedi-
ment from certain types of lakes, a geologist may look at the pollen
changes that have occurred through time and reconstruct the history of the
climate and vegetation in an area.
Volcanic ash thrown into the atmosphere during eruptions enters lakes
and forms a discrete layer of ash on the lake bottom. When Mount St. Hel-
ens erupted in 1980, it deposited several centimeters of ash in lakes more
than 160 kilometers east of the volcano. Geologists have used layers of ash
in lakes to reconstruct the history of volcanic eruptions in some areas. Al-
though dust storms contribute sediment to lakes, such storms are usually
too infrequent in most areas to contribute significant amounts.
Water Circulation
Lake waters are driven into circulation by temperature-induced density
changes and wind. Most freshwater lakes in temperate climates circulate
completely twice each year; they are termed dimictic lakes. Circulation ex-
erts a profound influence on water chemistry of the lake and the amount
and type of sediment present within the water column. During summer
stratification, the lake is thermally stratified into three zones. The upper
layer of warm water (epilimnion) floats above the denser cold water and
prevents wind-driven circulation from penetrating much below the
epilimnion. The epilimnion is usually in circulation, is rich in oxygen (from
algal photosynthesis and diffusion from the atmosphere), and is well
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Lakes and limnology
Riparian ecosystems vary widely, depending on the course of the river. Upper
courses, at higher and colder elevations, typically run faster and are highly oxy-
genated, supporting trout, bass, salmon, and other freshwater fish and their
predators, but not as many grasses, for example, as middle or lower courses and
estuaries. (PhotoDisc)
lighted. This layer is where summer blooms of green and blue-green algae
occur and calcite precipitation begins. The middle layer (thermocline) is a
transition zone in which the water cools downward at a rate of greater than
1 degree Celsius per meter. The bottom layer (hypolimnion) is cold, dark,
stagnant, and usually poor in oxygen. There, bacteria decompose the bot-
tom sediment and release phosphorus, manganese, iron, silica, and other
constituents into the hypolimnion.
Sediment deposited in summer includes a large amount of organic mat-
ter, clastic materials washed in during summer rainstorms, and endogenic
carbonate minerals produced within the lake. The most common carbon-
ate mineral is calcite (calcium carbonate). The regular deposition of calcite
in the summer is an example of cyclic sedimentation, a sedimentary event
that occurs at regular time intervals. This event occurs yearly in the sum-
mer season and takes place in the upper 2 or 3 meters of water. On satellite
photos, it is even possible to see the summer events as whitings on large
lakes, such as Lake Michigan.
As the sediment falls through the water column in summer, it passes
through the thermocline into the hypolimnion and onto the lake bottom.
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Lakes and limnology
As it sits on the bottom during the summer months, bacteria, particularly

anaerobic bacteria (those that thrive in oxygen-poor environments), begin
to decompose the organic matter. As this occurs, the dissolved carbon di-
oxide increases in the hypolimnion. If enough carbon dioxide is produced,
the hypolimnion becomes slightly acidic, and calcite and other carbonates
that fell to the bottom begin to dissolve. The acidic conditions also release
dissolved phosphorus, calcium, iron, and manganese into the hypolim-
nion, as well as some trace metals. Clastic minerals such as quartz, feld-
spar, and clay minerals are not affected in such brief seasonal processes,
but some silica from biogenic material such as diatom frustules can dis-
solve and enrich the hypolimnion in silica. As summer progresses, the
hypolimnion becomes more and more enriched in dissolved metals and
nutrients.
Autumn circulation begins when the water temperature cools and the
density of the epilimnion increases until it reaches the same temperature
and density as the deep water. Thereafter, there is no stratification to pre-
vent the wind from circulating the entire lake. When this happens, the
cold, stagnant hypolimnion, now rich in dissolved substances, is swept
into circulation with the rest of the lake water. The dissolved materials
from the hypolimnion are mixed into a well-oxygenated water column.
Iron and manganese that formerly were present in dissolved form now ox-
idize to form tiny solid particles of manganese oxides, iron oxides, and hy-
droxides. The sediment therefore becomes enriched in iron, manganese, or
both during the autumn overturn, the amount of enrichment depending
upon the amount of dissolved iron and manganese that accumulated dur-
ing summer in the hypolimnion. Dissolved silica is also swept from the
hypolimnion into the entire water column. In the upper water column,
where sunlight and dissolved silica become present in great abundance,
diatom blooms occur. The diatoms convert the dissolved silica into solid
opaline frustules.
As circulation proceeds, the currents may sweep over the lake bottom
and actually resuspend 1 centimeter or more of sediment from the bottom
and margins of the lake. The amount of resuspension that occurs each year
in freshwater lakes is primarily the result of the shape of the lake basin. A
lake that has a large surface area and is very shallow permits wind to keep
the lake in constant circulation over long periods of the year.
As winter stratification comes, an ice cover forms over the lake and pre-
vents any wind-induced circulation. Because the circulation is what keeps
the lake sediment in suspension, most sediment quickly falls to the bottom;
sedimentation then is minimal through the rest of winter. If light can pene-
trate the ice and snow, some algae and diatoms can utilize this weak light,
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Lakes and limnology
present in the layer of water just below the ice, to reproduce. Their settling
remains contribute small amounts of organic matter and diatom frustules.
At the lake bottom, the most dense water (that at 4 degrees Celsius) accu-
mulates. As in summer, some dissolved nutrients and metals can build up
in this deep layer, but because the bacteria that are active in releasing these
substances from the sediment are refrigerated, they work slowly, and not
as much dissolved material builds up in the bottom waters.
When spring circulation begins, the ice at the surface melts, and the lake
again goes into wind-driven circulation. Oxidation of iron and manganese
occurs (as in autumn), although the amounts of dissolved materials avail-
able are likely to be less in spring. Once again, nutrients such as phospho-
rus and silica are circulated out of the dark bottom waters and become
available to produce blooms of phytoplankton. Spring rains often hasten
the melting, and runoff from rain and snowmelt in the drainage basin
washes clastic materials into the lake. The period of spring thaw is likely to
be the time of year when the maximum amount of new allogenic (exter-
nally derived) sediment enters the lake.
Spring diatom blooms continue until summer stratification prevents
further replenishment of silica to the epilimnion. Thereafter, the diatoms
are succeeded by summer blooms of green algae, closely followed by
blooms of blue-green algae. Silica is usually the limiting nutrient for dia-
toms; phosphorus is the limiting nutrient for green and blue-green algae.
Diagenesis
After sediments are buried, changes occur; this process of change after
burial is termed diagenesis. Physical changes include compaction and
dewatering. Bacteria decompose much organic matter and produce gases
such as methane, hydrogen sulfide, and carbon dioxide. The “rotten-egg”
odor of black lake sediments, often noticed on boat anchors, is the odor of
hydrogen sulfide. After long periods of time, minerals such as quartz or
calcite slowly fill the pores remaining after compaction.
One of the first diagenetic minerals to form is pyrite (iron sulfide). Much
pyrite occurs in microscopic spherical bodies that look like raspberries;
these particles, called framboids, are probably formed by bacteria in areas
with low oxygen within a few weeks. In fact, the black color of some lake
muds and oozes results as much from iron sulfides as from organic matter.
Other diagenetic changes include the conversion of mineraloid particles
containing phosphorus into phosphate minerals such as vivianite and apa-
tite. Manganese oxides may be converted into manganese carbonates
(rhodochrosite). Freshwater manganese oxide nodules may form in high-
energy environments such as Grand Traverse Bay in Lake Michigan.
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Lakes and limnology
Lake Ecosystems
Freshwater and saline lakes account for 0.009 and 0.008 percent of the total
amount of water in the world, respectively. Although this is a minute frac-
tion of the world’s water—almost all of it is in the oceans and in glaciers—
lakes are an extremely valuable resource. In terms of ecosystems, lakes are
divided into a pelagial (open-water) zone and a littoral (shore) zone where
macrovegetation grows. Sediments free of vegetation that occur below the
pelagial zone are in the profundal zone.
The renewal times for freshwater and saline lakes range from 1 to 100
years and 10 to 1,000 years, respectively. The length of time varies directly
with lake volume and average depth, and indirectly with a lake’s rate of
discharge. The rate of renewal, or turnover time, for lakes is much less than
that of oceans and glacial ice, which is measured in thousands of years.
Eutrophication
The aging of a lake by biological enrichment is known as eutrophication.
The water in young lakes is cold and clear, with minimal amounts of plant
and animal life. The lake is then in the oligotrophic state. As time goes on,
streams that flow into the lake bring in nutrients such as nitrates and phos-
phates, which encourage aquatic plant growth. As the fertility in the lake
increases, the plant and animal life increases, and organic remains start ac-
cumulating on the bottom. The lake is now becoming eutrophic. Silt and
organic debris continue to accumulate over time, slowly making the lake
shallower. Marsh plants that thrive in shallow water start expanding and
gradually fill in the original lake basin. Eventually the lake becomes a bog
and then dry land.
This natural aging of a lake can take thousands of years, depending
upon the size of the lake, the local climate, and other factors. However, hu-
man activities can substantially accelerate the eutrophication process.
Among the problems caused by humans are the pollution of lakes by nutri-
ents from agricultural runoff and poorly treated wastewater from munici-
palities and industries. The nutrients encourage algal growth, which clogs
the lake and removes dissolved oxygen from the water. The oxygen is
needed for other forms of aquatic life. The lake has now entered a hyper-
eutrophic state as declining levels of dissolved oxygen result in incomplete
oxidation of plant remains, a situation that eventually causes the death of
the lake as a functioning aquatic ecosystem. In a real sense, the lake chokes
itself to death.
Research Methods
Scientists who study lakes (limnologists) must study all the natural
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Lakes and limnology
sciences—physics, chemistry, biology, meteorology, and geology—because

lakes are complex systems that include biological communities, changing
water chemistry, geological processes, and interaction among water, sun-
light, and the atmosphere. Many who study ecology become limnologists,
and vice versa.
Limnologists study modern lake sediments by collecting samples from
the water column in sediment traps (cylinders and funnels into which the
suspended sediment settles over periods of days or weeks) or by filtering
large quantities of lake water. Living material is often sampled with a
plankton net. Older sediments that have accumulated on the bottom are
collected with dredges and by piston coring, which involves pushing a
sharpened hollow tube (usually about 2.5 centimeters in diameter) down-
ward into the sediment. Cores are valuable because they preserve the sedi-
ment in the order in which it was deposited, from oldest at the bottom to
youngest at the top. Once the sample is collected, it is often frozen and
taken to the laboratory. There, pollen and organisms may be examined by
microscopy, minerals may be determined by X-ray diffraction, and chemi-
cal analyses may be made.
Varves are thin laminae that are deposited by cyclic processes. In fresh-
water lakes, each varve represents one year’s deposit; it consists of a cou-
plet with a dark layer of organic matter deposited in winter and a light-
colored layer of calcite deposited in summer. Varves are deposited in lakes
where annual circulations cannot resuspend bottom sediment and there-
fore cannot mix it to destroy the annual lamination. Some lakes that are
small and very deep may produce varved sediments; Elk Lake in Minne-
sota is an example. In other lakes, the accumulation of dissolved salts on
the bottom eventually produces a dense layer (monimolimnion), which
prevents disturbance of the bottom by circulation in the overlying fresher
waters. Soap Lake in Washington State is an example. Because each varve
couplet represents one year, a geologist may core the sediments from a
varved lake and count the couplets to determine the age of the sediment in
any part of the core. The pollen, the chemistry, the diatoms, and other con-
stituents may then be carefully examined to deduce what the lake was like
during a given time period. The study is much like solving a mystery from
a variety of clues. Eventually, the history of climate changes of the area
may be learned from the study of lake varves.
Edward B. Nuhfer and Robert Hordon
See also: Acid deposition; Ecosystems: definition and history; Eutrophica-

tion; Habitats and biomes; Marine biomes; Ocean pollution and oil spills;
Reefs; Wetlands.
372
Lakes and limnology

Bramwell, Martyn. Rivers and Lakes. London: Franklin Watts, 1986.
Burgis, Mary J., and Pat Morris. The Natural History of Lakes. New York:
Cole, Gerald A. Textbook of Limnology. 4th ed. Prospect Heights, Ill.: Wave-
land Press, 1994.
Fraser, Andrew S., Michael Meybeck, and Edwin D. Ongley. Water Quality
of World River Basins. 14th ed. New York: United Nations Publications,
1998.
Håkanson, Lars, and M. Jansson. Principles of Lake Sedimentology. New
York: Springer-Verlag, 1983.
Horne, Alexander J., and Charles R. Goldman. Limnology. 2d ed. New York:
McGraw-Hill, 1994.
Imberger, Jeorg, ed. Physical Processes in Lakes and Oceans. Washington,
D.C.: American Geophysical Union, 1998.
Lerman, Abraham, Dieter M. Imboden, and Joel R. Gat, eds. Physics and
Chemistry of Lakes. New York: Springer-Verlag, 1995.
Thornton, Kent W., Bruce L. Kimmel, and Forrest E. Payne, eds. Reservoir
Limnology: Ecological Perspectives. New York: John Wiley, 1990.
U.S. Environmental Protection Agency. The Great Lakes: An Environmental
Atlas and Resource Book. Chicago: Great Lakes Program Office, 1995.
Wetzel, Robert G. Limnology. 3d ed. San Diego: Academic Press, 2001.
373
LANDSCAPE ECOLOGY
Type of ecology: Landscape ecology
Humans live in natural landscapes that they have modified and managed to suit
their own needs of shelter, security, aesthetics, and usefulness. The science of man-
aging the habitat components of modified landscapes is called landscape ecology, a
burgeoning field concerned with preserving the naturalness of modified land-
scapes while minimizing the negative impact of human intrusion in natural habi-
tats within these landscapes.
K nowledge of landscape ecology is fundamentally important for a

number of reasons: natural resource planning, residential and com-
mercial development, wildlife conservation, forestry, agriculture, soil sci-
ence, ecological function, sociobiology, and the structure of urban and
suburban habitats are all elements of landscape ecology of continuing en-
vironmental and economic interest to humans.
Structurally, landscapes share four components: patches of habitat, cor-
ridors that connect patches of habitat, the background matrix that includes
patches and corridors, and the wildlife that inhabits the landscape. Hu-
mans and natural processes constantly modify each of these landscape
components.
From an ecological standpoint, humans most impact landscapes by
fragmenting large blocks of natural habitat into landscapes consisting of
patches of natural habitat, patches of modified habitat, and developed ar-
eas. For example, a forested landscape may be transformed into a mosaic of
woodland patches, farms, residences, parks, preserves, and greenbelts—
the whole typically crossed and gridworked by artificial corridors of road-
ways, power lines, and gas lines along with natural corridors such as
rivers, streams, and fence rows.
Some important management issues in landscape ecology include pre-
serving existing wildlife and wildlife habitats, maintaining and improving
wildlife habitat and biodiversity, minimizing detrimental impacts of habi-
tat fragmentation, creating natural corridors for wildlife movement be-
tween habitats, and creating and managing wildlife parks and preserves in
human modified landscapes.
Landscape Fragmentation
The splitting of a contiguous area of natural landscape into two or more
smaller blocks of habitat is called landscape fragmentation. The smaller
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Landscape ecology
habitat parcels that result are called fragments or patches. As human popu-
lations grow they appropriate more and more of the natural landscape to
accommodate their immediate needs for housing and agriculture, thus the
rate of landscape fragmentation continues to increase. Because human
populations are projected to grow for several more decades, at least, land-
scape fragmentation is considered to be a global wildlife and wildlife habi-
tat issue of immediate and serious concern.
Natural processes such as floods, ice storms, winds, and landslides con-
tribute to landscape fragmentation, but the vast majority of fragmentation
occurs through human activities. Logging, agriculture, roadways, rail
lines, power lines, gas lines, trails, the construction of houses, housing
clusters, commercial and industrial developments are all some of the ways
in which humans fragment natural habitats.
The Patchwork Mosaic

Habitat fragmentation ultimately results in a landscape mosaic compris-
ing a patchwork of artificial and natural habitats strewn across the land-
scape often in haphazard and unplanned fashion. Natural habitats, slightly
modified habitats, and totally altered habitats and the corridors that con-
nect them juxtaposition one another. The interrelationships of these natu-
ral and modified patches with respect to size, shape, and connectivity can
have profound impacts that collectively tend to limit natural habitats
within the landscape and the wildlife that occur in those natural habitats.
Some of the more important of these constraints include the decline or
elimination of habitat specialists or area-sensitive species, the invasion of
natural habitat patches by domestic, exotic, or alien species, the increased
mortality of natural species caused by vehicles or predation by pets and fe-
ral animals, and the disturbance of reproductive efforts and other natural
behaviors by the range of activities that invariably accompany human in-
trusion into and modification of, natural landscapes.
Initially, landscape fragmentation increases the number and variety of
habitat patches within a landscape, thereby increasing habitat diversity.
The many small patches provide “stepping stones” that promote species
dispersal and interchange from one patch to another which in turn pro-
motes gene flow between patches. Small patches may also provide habitat
for certain familiar species, such as the American robin (Turdus migratorius)
and gray catbird (Dumetella carolinensis), that use the edges of small patches
for foraging, nesting, or refuges. Also, a large number of small patches may
reduce erosion and ultimately loss of natural habitats that so often accom-
panies human activity and intrusion.
However, continued landscape fragmentation results in ever smaller
375
Landscape ecology
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habitat patches that necessarily support fewer species and smaller wildlife
populations, thereby increasing the risk of extinction of remaining species.
As habitat patches shrink in size or disappear, the remaining habitat
patches become increasingly isolated, inhibiting wildlife interchange be-
tween patches and making recolonization of the more remote habitat
patches increasingly difficult.
As habitat patches diminish in size, species populations begin to disap-
pear. The first to go are the species that need large areas for their activi-
ties such as predators that require large territories for their food base. Thus,
as patches shrink to a critical point the area cannot support sufficient
prey populations to sustain viable populations of larger predators, such
as northern goshawks (Accipiter gentilis) or red-shouldered hawk (Buto
lineatus).
Area-sensitive species and interior species are also increasingly placed
at risk. Area-sensitive species such as the Carolina chickadee (Poecile atri-
capilla) and eastern wood-peewee (Contopus virens) require large areas of
undisturbed habitat for nesting, foraging, and other activities. As habitat
patches decline in size, the numbers of these species dramatically decrease.
Interior species such as the ovenbird (Seiurus aurocapillus) select habitats
deep within the interior of natural habitats. As the size of habitat patches
decreases, there is insufficient amount of core area needed to sustain their
activities. The decline in populations of many neotropical migrants that
nest in woodlands of Eastern North America is attributed to habitat frag-
mentation.
376
Landscape ecology
The transformation of natural landscapes into human and natural

patches almost always creates a modified habitat suitable for intrusive spe-
cies that are tolerant of humans and human-modified landscapes, such as
the house sparrow (Passer domesticus), Eurasian starling (Sturnis vulgaris),
raccoon (Procyon lotor), and opossum (Didelphis virginiana), all of which
are behaviorally adapted to exploit the modified landscapes. Landscape
modification also provides an environment for pets that inevitably accom-
pany human intrusions, such as house cats (Felis catus) and dogs (Canis
familiaris), both of which may predate natural wildlife species. Both intru-
sive and domesticated species reduce biodiversity either through outright
predation or by competition.
A critical concern to landscape ecologists involves the ratio of edge hab-
itat to interior habitat in habitat patches resulting from fragmentation. As
habitat patches get smaller, the ratio of edge habitat to interior habitat of
each patch increases. Greater edge habitat provides favorable habitat for
game and other edge species, but interior species and area-sensitive spe-
cies are placed at increased risk because predators, scavengers, and para-
sitic species can now penetrate farther into the interior of the habitat patch.
Some of the more familiar and important avian edge species in terms of
their impacts on interior species include the brown-head cowbird (Molo-
thrus ater) and the blue jay (Cyanocitta cristata). Two of the many species of
mammals that frequent edge habitat and may become nuisance species in
human-modified landscapes include the raccoon and the opossum.
The potential impact of edge species on interior species is exemplified
by a small blackbird with a distinctive brown head called the brown-
headed cowbird. The cowbird is a brood parasite that frequents open
woodland, grasslands, and forest edge. Unlike other birds, the cowbird
does not build a nest but rather deposits its eggs in the nests of other birds,
especially flycatchers, vireos, warblers, orioles, and finches. Female brown-
headed cowbirds visit nests of potential host species during construction.
Once the host bird lays its clutch, the female cowbird visits the nest and re-
moves one or more of the eggs, either by eating them or by dropping them
over the edge of the nest cup. She always leaves one egg—otherwise the
host might not return. She then deposits a single egg in the host nest. The
host female returns and incubates the clutch, which now includes the cow-
bird egg. The cowbird egg hatches a day or two before the other eggs in the
nest and the young grows quickly, demanding and receiving a large por-
tion of the food that the host adult birds bring to the nestlings. The impact
of cowbird brood parasitism increases with increasing landscape fragmen-
tation. As patches get smaller the ratio of edge habitat to core habitat in a
given habitat patch increases, exposing more of the interior to cowbird ac-
377
Landscape ecology
tivity. Cowbirds penetrate deeper into habitats to parasitize more nests of

interior species. In fact, increased cowbird brood parasitism accompany-
ing habitat fragmentation is one reason that interior woodland species
such as wood warblers are thought to be declining.
Corridors and Connectivity

Corridors are links of natural habitats that connect landscape patches to
one another. Examples of corridors include areas of vegetation left unde-
veloped by human disturbance such as belts of vegetation along rivers and
streams, vegetated strips along roadways, railways, and drainage ditches,
vegetation strips beneath power lines and above gas lines, and linear ex-
tensions of vegetated areas such as windbreaks, fence rows, and hedge-
rows.
Corridors are critically important elements of landscape ecology that
help maintain stability and diversity of wildlife within each of the natural
patches by providing dispersal routes for plants and animals between hab-
itat patches. The ability of wildlife to use these corridors to disperse from
one habitat patch to another is called connectivity. Larger patches that sup-
port larger species populations serve as the source of individuals that dis-
perse along corridors to repopulate or augment species populations in
smaller patches, where species populations are low, are declining in num-
bers, or have been extirpated.
Corridors of natural vegetation promote movement of organisms from
one habitat patch by reducing their vulnerability during dispersal. Corri-
dors also facilitate gene flow between wildlife populations in each patch,
reducing the change of inbreeding depression. Conversely, lack of corri-
dors or low corridor connectivity puts dispersing wildlife at greater risk.
For example, if vegetated corridors are lacking woodland, animals must
cross open areas to disperse from one woodland habitat patch to another,
increasing their vulnerability to predators. Some animals, such as certain
amphibians and reptiles, may be unable to disperse from one habitat patch
to another if connecting corridors are not available.
Corridor width is also an important consideration in landscape ecology.
Narrow corridors increase the potential for intrusion of domestic animals
such as cats and dogs along and into the corridor, making wildlife dis-
persal more difficult and more dangerous. Narrow corridors may also con-
tribute to poaching of larger animals, which are more visible and therefore
more vulnerable during their dispersal within the confines of narrow corri-
dors. However, low connectivity also decreases the rate of spread of inva-
sive species and pests, reduces the dispersal of pollutants, thereby enhanc-
ing survival of interior species that occupy patches.
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Landscape ecology
Some types of natural corridors also have distinct ecological character-

istics that contribute to the overall species and landscape biodiversity in a
given landscape. For example, belts of vegetation along rivers and streams
form a distinctive ecological habitat called a riparian community that is in-
habited by riparian species as well as edge and dispersal species.
Parks and Preserves as Landscape Components

Parks and preserves are natural units of landscape that have been deliber-
ately set aside as protected parcels in an attempt to maintain ecologically
functioning communities of plants and animals. Their size is usually a con-
sequence of how valuable the land is that is being set aside, the value that
humans place on wildlife and wildlife communities, the types of land that
can be set aside, and the economic sacrifices that humans are willing to
make to ensure that natural wildlife communities are protected.
Much of the debate over size of parks and preserves has focused on
whether to establish a single large preserve or several small preserves. This
debate is sometimes called the SLOSS controversy, the acronym standing
for “single larger or several smaller” preserves.
Large preserves are favored by many ecologists because they provide
greater amounts of natural habitat that can provide the resource base
needed to support greater wildlife diversity and also larger populations of
each plant and animal species. A larger species population reduces the risk
of extinction or extirpation due to chance events such as skewed sex ratios
or poor reproductive rates in a given year or set of years. A major draw-
back of a single large preserve, however, is that all of the wildlife of an area
is concentrated within a limited landscape that can be destroyed by a sin-
gle catastrophe such as a severe rainstorm, floods, fires, ice storms, heavy
snowfall, landslides, and other natural catastrophes. Furthermore, concen-
tration of wildlife populations in a single large preserve exposes all of the
members of a species population to risk from pollutants, chance appear-
ance of competitors, or invasion of exotic species.
The alternate choice of creating several small preserves distributes
members of a species population among several preserves rather than
concentrating them all in a single large preserve. This greatly reduces the
chance that a catastrophe will eliminate the wildlife population, as it is ex-
tremely unlikely that a chance catastrophe will impact all of the smaller
preserves simultaneously. However, the smaller resource base of a smaller
preserve automatically limits numbers of individuals in each species pop-
ulation, thereby greatly increasing risk of extirpation of a given species
population in a smaller preserve due to chance events. For example, small
populations face increased risks of genetic inbreeding, the chance accumu-
379
Landscape ecology
lation of deleterious mutations, and genetic drift. Environmentally, smaller

species populations are also more susceptible to disease, the introduction
of predators, and the possibility of a skewed sex ratio that results in low re-
production in a given generation. To compensate for the increased risk of
wildlife extinction in smaller preserves, landscape ecologists must provide
corridors to connect preserves with one another, thereby promoting wild-
life movement between preserves.
Managing and maintaining parks and preserves and their connecting
corridors depends on several factors. Virtually all parks and preserves in
urban and suburban landscapes require developing and implementing
management strategies to ensure that disturbances from adjacent human-
modified habitats is minimized. Rare habitats or habitats that contain rare
or endangered wildlife must be managed differently from open-space par-
cels that contain common and widespread species. In some locales, parks
and preserves may best be managed under the multiple-use concept, func-
tioning simultaneously as wildlife preserves and as areas that provide rec-
reational opportunities for hiking, biking, bird-watching, and wildlife ap-
preciation.
Dwight G. Smith

Forman, Richard T. T., and M. Godron. Landscape Ecology. New York: John
Wiley & Sons, 1986.
Forman, Richard T. T. Land Mosaics: The Ecology of Landscapes and Regions.
Gardner, R. H., R. V. O’Neill, and M. G. Turner. Landscape Ecology in Theory
and Practice. New York: Springer, 2001.
Garner, H. F. The Origin of Landscapes: A Synthesis of Geomorphology. New
Gergel, S. E., and Monica G. Turner, eds. Learning Landscape Ecology: A Prac-
tical Guide to Concepts and Techniques. New York: Springer, 2001.
Gutzwiller, K. J. Applying Landscape Ecology in Biological Conservation. New
York: Springer, 2002.
Jongman, R. H. G., C. J. F. Ter Braak, and P. F. R. Van Tongeren. Data Analy-
sis in Communities and Landscape Ecology. New York: Cambridge Univer-
sity Press, 1995.
380
LICHENS
Lichens are an example of a very specialized “ecosystem” composed of two distinct

species, a fungus and a photosynthetic alga (or bacterium) that have coevolved to
live in a symbiotic relationship with each other in a community that grows on
rocks, trees, and other substrates.
L ichens are classified as members of the kingdom Fungi, with most be-
ing placed under the phyla Ascomycota and Basidiomycota. It is esti-
mated there are seventeen thousand species of lichen, representatives of
which have been found nearly everywhere in the world.
Symbiosis
Symbiosis is an extreme form of an ecological relationship or mutualism
between members of different species, in which each partner in the union
derives benefits from the other. In symbiotic unions, the partners are so de-
pendent on each other they can no longer independently survive.
In lichens, the fungal (mycobiont) symbiont provides protection, while
the green-algal or cyanobacterial (photobiont) symbiont provides sugars,
created by photosynthesis. It is often suggested that the fungus in lichen
species might also pass water and nutrients to the photobiont, but this
function is less well documented. This special relationship allows lichens
to survive in many environments, such as hot deserts and frozen Arctic
tundra, that are inhospitable to most other life-forms. As a result, the lichen
whole is greater than the sum of its parts. While in nature lichen partners
always exist together, under laboratory conditions it is possible to take the
lichen apart and grow the two partners separately.
Anatomy
Whereas in most plant species the anatomy of the organism is identified
with structures associated with a single vegetative body, the “lichen body”
is more aptly described as a colony of cells that share a variety of associa-
tions with one another that vary from one species of lichen to the next. In
some species of lichen, fungal and algal cells merely coexist. Coenogonium
leprieurii, for example, is a lichen that lives in low-light tropical and sub-
tropical forests in which the filamentous green-algal partner (Trebouxia) is
dominant.
381
Lichens
In most lichen species, however, the relationship between the symbiotic

partners is more intimate, with the lichen body appearing to be a single en-
tity. In these species the algal symbiont has no cell walls and is penetrated
by filaments, or haustoria, from the fungal symbiont. The haustoria pass
sugars from the algal cell to the fungal cell and may have a role in the trans-
portation of water and nutrients from the fungal cell back to the algal cell.
This integration is so complete that many naturalists prior to the nine-
teenth century mistakenly classified lichens as mosses.
In most lichen species it is nevertheless possible, with a good magnify-
ing device, to identify several distinct regions of the thallus or lichen body.
The outermost region is the cortex, a compacted layer composed of short,
thick hyphae (widely dilated filaments) of the fungal symbionts that pro-
tect the lichen from abiotic factors in its environment. These hyphae extend
downward into a second region, the photobiont layer, where they sur-
round the algal symbionts. Below this is a third region, the medulla, com-
posed of a loosely woven network of hyphae.
Underneath this is a fourth region, the undercortex, that is similar in ap-
pearance and structure to the cortex. The bottom of the lichen body is com-
Image Not Available
382
Lichens
posed of rhizines, rootlike structures composed of bundles of hyphae that

attach the lichen to its substrate (the rock, bark, or other support on which
it resides). This arrangement of regions into layers serves to prevent water
loss. Many species can survive complete desiccation, coming back to life
when water becomes available again. The cortex also contains pseudo-
cyphellae, which are pores that allow for the exchange of gases necessary
for photosynthesis.
Life Cycle
Lichens typically live for ten years or more, and in some species the lichen
body can survive for more than a hundred years. Reproduction in most
fungal species proceeds by the development of a cup- or saucer-shaped
fruiting body called an apothecium, which releases fungal spores to its sur-
rounding. Procreation in lichens is more problematic, in that the fungal off-
spring must also receive the right algal symbiont if they are to survive. The
most common form of dispersion in lichen is by the accidental breaking off
of small pieces of the thallus called isidia, which are then spread by wind to
new substrates. In some species, small outgrowths of the thallus known as
soralia arise, composed of both fungi and algae and surrounded by
hyphae, to form soredia, which after dispersion give rise to a new thallus.
Ecological and Economic Importance

Lichens not only demonstrate some basic ecological concepts, such as
mutualism and symbiosis, but also are excellent bioindicators of air pollu-
tion, as many species are particularly sensitive to certain contaminants in
their surroundings, such as sulfur dioxide. They also play an important
role in tundra biomes, functioning as a major source of food for reindeer
and cattle in Lapland. One species of lichen (Umbilicaria esculenta) is con-
sidered a delicacy in Japan. Historically, lichens have been used as pig-
ments for the dying of wool. The medical properties of some species of li-
chens for lung disease and rabies have led to a renewed interest in them.
David W. Rudge
See also: Animal-plant interactions; Coevolution; Communities: ecosys-

tem interactions; Communities: structure; Convergence and divergence;
Food chains and webs; Mycorrhizae; Symbiosis; Trophic levels and ecolog-
ical niches.

Brodo, Irwin M., Sylvia Duran Sharnoff, and Stephen Sharnoff. Lichens of
North America. New Haven, Conn.: Yale University Press, 2001.
383
Lichens
Dobson, Frank. Lichens: An Illustrated Guide. Richmond, Surrey, England:

Richmond, 1981.
Hale, Mason E., Jr. The Biology of Lichens. 3d ed. London: Edward Arnold,
1983.
Hawksworth, D. L., and D. J. Hill. The Lichen-Forming Fungi. New York:
Blackie & Son, 1984.
Purvis, William. Lichens. Washington, D.C.: Smithsonian Institution Press,
2000.
384
MAMMALIAN SOCIAL SYSTEMS
Social organization in mammals ranges from solitary species, which come together
only to breed, to large and intricately organized societies. Understanding the social
systems of mammals is essential for effective conservation of species.
A ll levels of social organization occur in mammals. There are solitary

species, such as the mountain lion (Felis concolor), in which the male
and female adults come together only to mate and the female remains with
her young only until they are capable of living independently. At the other
numerical extreme are some of the hoofed mammals, which form herds of
thousands of individuals. The most socially specialized mammal is proba-
bly Africa’s naked mole rat (Heterocephalus glaber), which has a eusocial col-
ony structure similar to that of ants, bees, and termites. No current theory
accounts for the diversity of mammalian social systems, but two broad
generalizations are consistently employed to explain mammalian species’
social organization. These are the environmental context in which the spe-
cies exists and the mammalian mode of reproduction.
Reproductive Determinants
More than any other group of animals, mammals are required to form
groups for at least part of their lives. Although in all sexually reproducing
animals the sexes must come together to mate, mammals have an addi-
tional required association between mother and young: All species of
mammal feed their young with milk from the mother’s mammary glands.
This group, a female and her young, is the basis for the development of
mammalian social groups. In some species, the social group includes sev-
eral females and their young and may involve one or more males as well.
Environmental Determinants
Mammalian societies are always organized around one or more females
and their offspring. Males may also be part of the group, or they may form
separate groups. The size and structure of the group are determined by the
ecological setting in which it evolves. The particular ecological factors that
seem to be of greatest importance in this determination are food supply,
the distribution of the food, and predation (including the hiding places
and escape routes available in the habitat).
Large groups occur when food is scattered in a patchy distribution.
385
Mammalian social systems
These groups are largest when the patches contain abundant food. Many
organisms are more likely to find the scattered patches than is a single indi-
vidual. As long as the patches have enough food for all members of the
group, it is to each member’s advantage to search with the group. On the
other hand, if food is evenly dispersed in small units throughout the envi-
ronment, the advantage of a group search is lost. Each individual will be
better off searching for itself, and some strategy involving a very small so-
cial group or even solitary existence would be advantageous.
A somewhat similar argument follows for predators. If large prey are
taken, a group of predators should be able to subdue the prey and protect
its remains from scavengers more efficiently. If small prey are taken, soli-
tary predators have the advantage, since the prey is easily dispatched and
the predator will have it to itself. Many other factors are involved in deter-
mining the final form of a species’ social organization, but the family unit
and environmental context are fundamental in determination of all mam-
malian social structures.
Primate Social Organization

The primates are the most social group of mammals. Monkeys demon-
strate the importance of food supply and its distribution in determining
social structure. The olive baboon (Papio anubis) occupies savannas, where
Lions are among the few feline

species that live in groups,
called prides. Each pride
consists of only one or a few
males and several or many
females. The females share the
responsibility for raising the
cubs, teaching them hunting
behaviors and other survival
skills. These four littermates
are nearly independent and, if
male, will likely leave the
pride. (PhotoDisc)
386
it exists in large groups of several adult males, several adult females, and
their young. Finding fifty or more animals in a group is not uncommon. In-
dividual males do not guard or try to control specific females except when
the females are sexually receptive. The group’s food supply is in scattered
patches, but each patch contains an abundance of food. The advantage of
having many individuals searching for the scattered food is obvious: If any
member finds a food-rich patch, there is plenty for all.
Predation probably also plays a role in the olive baboon’s social organi-
zation. The savannas they roam have many predators and few refuges for
escape. A large group is one defense against predators if hiding or climb-
ing out of reach is not practical. Having many observers increases the
chance of early detection, giving the prey time to elude the predator. A
large group can also mount a more effective defense against a predator.
Large groups of baboons use both of these tactics.
The hamadryas baboon (Papio hamadryas), on the other hand, lives in
deserts in which the food supply is not only scattered but also often found
in small patches. The hamadryas baboon’s social structure contrasts with
that of the olive baboon, perhaps because the small patches do not supply
enough food to support large groups. A single adult male, one or a few
adult females, and their young make up the basic group of fewer than
twenty individuals. Several of these family groups travel together under
certain conditions, forming a band of up to sixty animals. Within the band,
however, the family groups remain intact. The male of each group herds
his females, punishing them if they do not follow him. The bands are prob-
ably formed in defense against predators. They break up into family units
if predators are absent. At night, hamadryas baboons sleep on cliffs, where
they are less accessible to predators. Because suitable cliffs are limited,
many family groups gather at these sites. Hundreds of animals may be in
the sleeping troop, probably affording further protection against preda-
tors.
Though there are exceptions, forest primates consistently live in smaller
groups. In many species, fewer than twenty individuals make up the social
group at all times. These consist of one or a few mature males, one or a few
mature females, and their offspring. The groups are more evenly distrib-
uted throughout their habitat than are groups of savanna or desert pri-
mates. In forests, the food supply is more abundant and more evenly dis-
tributed. Escape from predators is also more readily accomplished—by
climbing trees or hiding in the dense cover. Under these conditions, the ad-
vantages of large groups are minimal and their disadvantages become ap-
parent. For example, in small groups the competition for mates and food is
less.
387
Ungulate Social Organization

The ungulates have all levels of social organization. African antelope dem-
onstrate social organizations that, in some ways, parallel those of the pri-
mates. Forest antelope such as the dik-dik (Madoqua) and duiker (Ceph-
alophus) are solitary or form small family groups, and they are evenly
spaced through their environment. Many hold permanent territories con-
taining the needs of the individual or group. They escape predators by hid-
ing and are browsers, feeding on the leaves and twigs of trees.
Many grassland and savanna antelope, such as wildebeest (Conno-
chaetes), on the other hand, occur in large herds. They outrun or present a
group defense to predators and are grazers, eating the abundant grasses of
their habitat. In many cases, they are also migratory, following the rains
about the grasslands to find sufficient food. The social unit is a group of re-
lated females and their young. Males leave the group of females and young
as they mature. They join a bachelor herd until fully mature, at which time
they become solitary, and some establish territories. The large migratory
herds are composed of many female-young groups, bachelor herds, and
mature males. The social units are maintained in the herd. Though it may
seem strange to speak of solitary males in a herd of thousands, that is their
social condition. The male territories are permanent in areas that have a re-
liable food supply year-round, but they cannot be in regions in which
the species is migratory. Under these conditions, the males set up tempo-
rary breeding territories wherever the herd is located during the breeding
season.
There are parallels with primate social patterns. Large groups are
formed in grasslands, and these roam widely in search of suitable food.
The groups are effective as protection against predators in habitats with
few hiding places. Smaller groups are found in forests, where food is more
evenly dispersed and places to hide from predators are more readily
found.
Rodent Social Organization

Rodents also have all kinds of social organization. The best known, and
one of the most complex, is the social system of the black-tailed prairie dog
(Cynomys ludovicianus). The coterie is the family unit in this case, and it
consists of an adult male, several adult females, and their young. Members
maintain a group territory defended against members of other coteries.
Coterie members maintain and share a burrow system. Elaborate greeting
rituals have developed to allow the prairie dogs of a coterie to recognize
one another. Hundreds of these coteries occur together in a town. The
members of these towns keep the vegetation clipped—as a result, preda-
388
tors can be seen from a distance. Prairie dogs warn one another with a
“bark” when they observe a predator, and the burrow system affords a ref-
uge from most predators.
The only vertebrate known to be eusocial is the naked mole rat. It occurs
in hot, dry regions of Africa. The colony has a single reproductive female, a
group of workers, and a group of males whose only function is to breed
with the reproductive female. The workers cooperate in an energetically
efficient burrowing chain when enlarging the burrow system. In this way,
they are able to extend the burrow system quickly during the brief wet sea-
son. Digging is very difficult at other times of the year. The entire social
system is thought to be an adaptation to a harsh environment and a sparse
food supply.
Carnivore Social Organization

Most carnivores are not particularly social, but some do have elaborate so-
cial organization. Many of these are based on the efficiency of group hunt-
ing in the pursuit of large prey or on the ability of a group to defend a large
food supply from scavengers. The gray wolf (Canis lupus) and African
hunting dog (Lycaon pictus) are examples. In both cases, the social group, or
pack, consists of a male and female pair and their offspring of several
years. Though there are exceptions, solitary carnivores and carnivores that
form temporary family units during the breeding season, such as the red
fox (Vulpes vulpes), hunt prey smaller than themselves. The coyote (Canis
latrans) can switch social systems to use the food available most efficiently.
It forms packs similar to those of the gray wolf when its main prey is large
or when it can scavenge large animals and is solitary when the primary
available prey is small.
These examples and many others show that the social groups of mam-
mals are based on the family group. The particular social organization em-
ployed by a species is determined by the ecological situation in which it oc-
curs. The specific aspects of the environment that seem to be most
important include food abundance, food distribution, food type, and pro-
tection from predators.
Carl W. Hoagstrom
See also: Altruism; Communication; Defense mechanisms; Displays; Ethol-

ogy; Habituation and sensitization; Herbivores; Hierarchies; Insect societ-
ies; Isolating mechanisms; Migration; Mimicry; Omnivores; Pheromones;
Poisonous animals; Predation; Reproductive strategies; Territoriality and
aggression.
389

Dunbar, Robin I. M. Primate Social Systems. Ithaca, N.Y.: Cornell University
Press, 1988.
Eisenberg, John F., and Devra G. Kleiman, eds. Advances in the Study of
Mammalian Behavior. Special Publication 7. Shippensburg, Pa.: Ameri-
can Society of Mammalogists, 1983.
Gittleman, John L., ed. Carnivore Behavior, Ecology, and Evolution. Ithaca,
N.Y.: Cornell University Press, 1989.
Immelmann, Klaus, ed. Grzimek’s Encyclopedia of Ethology. New York: Van
Nostrand Reinhold, 1977.
Macdonald, David W. European Mammals: Evolution and Behavior. London:
HarperCollins, 1995.
Nowak, Ronald M., and John L. Paradiso. Walker’s Mammals of the World.
6th ed. 2 vols. Baltimore: Johns Hopkins University Press, 1999.
Rosenblatt, Jay S., and Charles T. Snowdon, eds. Parental Care: Evolution,
Mechanisms, and Adaptive Significance. Advances in the Study of Behav-
ior 25. San Diego, Calif.: Academic Press, 1996.
Slater, P. J. B. An Introduction to Ethology. Reprint. London: Cambridge Uni-
Vaughan, Terry A. Mammalogy. 4th ed. Philadelphia: Saunders College
Publishing, 2000.
Wrangham, Richard W., W. C. McGrew, Frans B. M. De Waal, and Paul G.
Heltne, eds. Chimpanzee Cultures. Cambridge, Mass.: Harvard Univer-
sity Press, 1994.
390
MARINE BIOMES
ecology
The world’s oceans contain the largest and most varied array of life-forms on earth.
The marine environment is divided into coastal, open water, deep-sea, and bottom
zones and the lives of animals living in each of these regions are dictated by the
physical conditions present in these zones.
A pproximately 71 percent of earth’s surface is covered by salt water,

and the marine environments contained therein constitute the largest
and most diverse array of life on the planet. Life originated in the oceans,
and the salt water that comprises the largest constituent of the tissues of all
living organisms is a vestigial reminder of the aquatic origins of life.
Marine Zones
The marine environment can be divided broadly into different zones, each
of which supports numerous habitats. The coastal area between the high
and low tide boundaries is known as the intertidal zone; beyond this is the
neritic zone, relatively shallow water that extends over the continental
shelves. The much deeper water that extends past the boundaries of the
continental shelves is known as the oceanic zone. Open water of any depth
away from the coastline is also known as the pelagic zone. The benthic
zone is composed of the sediments occurring at the sea floor. Areas in
which freshwater rivers empty into the saltwater oceans produce a contin-
ually mixed brackish water region known as an estuary. Estuarine zones
often also include extensive wetland areas such as mudflats or salt
marshes.
Zones in the marine environment are distributed vertically as well as
horizontally. Life in the ocean, as on land, is ultimately supported by sun-
light in most cases, used by photosynthetic plants as an energy source.
Sunlight can only penetrate water to a limited depth, generally between
one hundred and two hundred meters; this region is known as the photic
or epipelagic zone. Below two hundred meters, there may be sufficient
sunlight penetrating to permit vision, but not enough to support photo-
synthesis; this transitional region may extend to depths of one thousand
meters and is known as the disphotic or mesopelagic zone. Below this
depth, in the aphotic zone, sunlight cannot penetrate and the environment
is perpetually dark, with the exception of small amounts of light produced
391
Marine biomes
by photoluminescent invertebrate and vertebrate animals. This aphotic

zone is typically divided into the bathypelagic zone, between seven hun-
dred and one thousand meters as the upper range and two thousand to
four thousand meters as the lower range, where the water temperature is
between 4 and 10 degrees Celsius. Beneath the bathypelagic zone, overly-
ing the great plains of the ocean basins, is the abyssalpelagic zone, with a
lower boundary of approximately six thousand meters. Finally, the deep-
est waters of the oceanic trenches, which extend to depths of ten thousand
meters, constitute the hadalpelagic zone. In each of these zones, the nature
and variety of marine life present is dictated by the physical characteristics
of the zone. However, these zones are not absolute, but rather merge grad-
ually into each other, and organisms may move back and forth between
zones.
Plankton
Marine life can be divided broadly into three major categories. Those small
organisms that are either free-floating or weakly swimming and which
thus drift with oceanic currents are referred to as plankton. Plankton can
be further divided into phytoplankton, which are plantlike and capable
of photosynthesis; zooplankton, which are animal-like; and bacterio-
plankton, which are bacteria and bluegreen algae suspended in the water
column. Larger organisms that can swim more powerfully and which can
thus move independently of water movements are known collectively as
the nekton. Finally, organisms that are restricted to living on or in the sedi-
ments of the seafloor bottom are referred to as the benthos.
The phytoplankton, which are necessarily restricted to the photic zone,
are by far the largest contributors to photosynthesis in the oceans. The
phytoplankton are therefore responsible for trapping most of the solar en-
ergy obtained by the ocean (the primary productivity), which can then be
transferred to other organisms when the phytoplankton are themselves
ingested. The phytoplankton are composed of numerous different types
of photosynthetic organisms, including diatoms, which are each encased
in a unique “pillbox” shell of transparent silica, and dinoflagellates. The
very rapid growth of some species of dinoflagellates in some areas results
in massive concentrations or blooms that are sometimes referred to as
red tides. Chemicals that are produced by red tide dinoflagellates often
prove toxic to other marine organisms and can result in massive die-offs of
marine life. Smaller photosynthetic plankton forms comprise the nano-
plankton and also play an important role in the photosynthetic harnessing
of energy in the oceans.
The zooplankton are an extremely diverse group of small animal organ-
392
Marine biomes
isms. Unlike the phytoplankton, which can make their own complex or-
ganic compounds via photosynthesis, the phytoplankton must ingest or
absorb organic compounds produced by other organisms. This is accom-
plished by either preying upon other planktonic organisms or by feeding
on the decaying remains of dead organisms. A number of zooplankton
species also exist as parasites during some portion of their life cycles, living
in or upon the bodies of nekton species. The largest group of zooplankton
are members of the subphylum Crustacea, especially the copepods. These
organisms typically possess a jointed exoskeleton, or shell, made of chitin,
large antennae, and a number of jointed appendages. Space precludes a
definitive listing of all of the zooplanktonic organisms, however virtually
all of the other groups of aquatic invertebrates are represented in the bewil-
dering variety of the zooplankton, either in larval or adult forms. Even
fish, normally a part of the nekton, contribute to the zooplankton, both as
eggs and as larval forms.
The bacterioplankton are found in all of the world’s oceans. Some of
these, the blue-green algae (cyanobacteria), play an important role in the
photosynthetic productivity of the ocean. Bacterioplankton are usually
found in greatest concentrations in surface waters, often in association
with organic fragments known as particulate organic carbon, or marine
snow. Bacterioplankton play an important role in renewing nutrients in the
photic zones of the ocean; such renewal is important in maintaining the
photosynthetic activity of the phytoplankton, upon which the rest of ma-
rine life is in turn dependent.
One of the principal problems facing plankton is maintaining their posi-
tion in the water column. Since these organisms are slightly denser than
the surrounding seawater, they tend to sink. Clearly this is a disadvantage,
particularly since plankton typically have very limited mobility. This is es-
pecially true for the photosynthetic phytoplankton, which must remain
within the photic zone in order to carry on photosynthesis. A number of
strategies have evolved among planktonic species to oppose this tendency
to sink. Long, spindly extensions of the body provide resistance to the flow
of water. Inclusions of oils or fats (which are less dense than water) within
the body provide positive buoyancy by decreasing the overall density of
the plankton. Finally, some species, such as the Portuguese man-o’-war,
generate balloonlike gas bladders, which provide enough buoyancy to
keep them at the very surface of the epipelagic zone.
Nekton
The nekton comprise those larger animals that have developed locomotion
to a sufficient degree that they can move independently of the ocean’s
393
Marine biomes
water movements. Whereas the plankton are principally invertebrates,

most of the nekton are vertebrates. The majority of the nekton are fish, al-
though reptile, bird, and mammalian species are also constituent parts.
The oceanic nekton are those species which are found in the epipelagic
zone of the open ocean. These include a wide variety of sharks, rays, bony
fish, seabirds, marine mammals, and a few species of reptiles. Some mem-
bers of the oceanic nekton, such as blue sharks, oceanic whitetip sharks,
tuna, flying fish, and swordfish, spend their entire lives in the pelagic envi-
ronment; these are said to be holoepipelagic. Others, the meroepipelagic
nekton—such as herring, dolphins, salmon, and sturgeon—spend only a
portion of their lives in the epipelagic zone, returning to coastal or fresh-
water areas to mate.
Seabirds are a special case: Although they spend much of their time fly-
ing over the epipelagic zone and nest on land, they feed in the epipelagic
zone. Some species may dive as deep as one hundred meters in search of
prey. Some members of the nekton enter the epipelagic only at certain
times in their life cycles. Eels of the family Anguillidae spend most of their
lives in fresh water but return to the epipelagic zone to spawn. Addi-
tionally, at night many species of deep-water fish migrate up into the
epipelagic to feed before returning to deeper waters during the daylight
hours.
The pelagic environment, unlike the terrestrial one, is profoundly three-
dimensional. Nektonic animals can move both horizontally and vertically
within the water column. Furthermore, since most of the pelagic environ-
ment is essentially bottomless, since there is no apparent or visible ground
or substrate, the environment is essentially uniform and featureless. These
features play an important role in the evolution of the behavior of nektonic
animals. Fish suspended in an essentially transparent and featureless me-
dium have no shelter in which to hide from predators, nor are there any ap-
parent landmarks to serve as directional cues for animals moving horizon-
tally from place to place. Life in the open ocean has therefore favored
adaptations for great mobility and speed with which to move across large
distances and escape from predators, as well as camouflage and cryptic
coloration designed to deceive potential predators or prey.
As is the case for plankton, most nektonic animals are denser than the
surrounding seawater, and maintaining position in the water column is of
the first importance. Most fish possess a swim bladder, a gas-filled mem-
branous sac within their body that opposes the tendency to sink and pro-
vides the fish with neutral buoyancy. Sharks and rays lack a swim bladder,
but accumulate large concentrations of fats and oils in their liver, which
also help counter the tendency to sink. Large, fast-swimming species of
394
Marine biomes
Seabirds and fish are

examples of nekton,
generally vertebrates that
have developed locomotion
to a sufficient degree that
they can move
independently of the ocean’s
water movements or occupy
other portions of the photic,
or epipelagic, zone of the
ocean ecosystem.
(PhotoDisc)
shark, tuna, and many billfish also rely on the generation of hydrodynamic
lift to maintain vertical position in the water column. The tail and body of
these fish generate forward thrust, moving the animal through the water,
and the fins, notably the pectoral fins, generate lift from the water flowing
over them in a manner similar to that of an airplane’s wing. Thus these ani-
mals fly through the water, but are in turn required to move continuously
in order to generate lift.
All members of the nekton are carnivores, feeding on other nektonic
species or upon plankton, particularly the larger zooplankton. In general,
the size of the prey consumed by nekton is directly related to the size of the
predator, with larger species consuming larger prey. However, the organ-
isms that feed upon plankton, the planktivores, include a wide variety of
fish species such as herring, salmon, and the whale shark, the largest extant
fish species. They also include the largest marine animals of all, the baleen
whales. The case of large animals feeding upon very small plankton di-
rectly addresses the need of all animals to meet their energy requirements.
For all animals, the amount of energy obtained from food consumed must
necessarily exceed the energy expended in acquiring the prey. Very large
animals, such as whales and whale sharks, require a great deal of energy to
move their bodies through the aquatic environment, but because of their
395
Marine biomes
great size they are necessarily less agile than smaller forms. The amount of
energy required to chase and catch these smaller animals would generally
exceed the energy derived from ingesting them. Plankton, however, are
relatively easy to obtain due to their very limited mobility. However, be-
cause of their small size, vast quantities of plankton must be ingested in or-
der to meet the metabolic requirements of large marine animals. Some very
large species that are not planktivores solve the energy problem by evolv-
ing behaviors for acquiring specialized diets that yield higher energy.
White sharks, for example, feed on fish when young, but as they age and
increase in size, marine mammals, notably seals and sea lions (pinnipeds),
become a major part of their diet. Marine mammals all possess blubber, an
energy-rich substance that yields much more energy than fish. Similarly,
sperm whales, the largest hunting carnivores on the planet, have a diet that
consists in large part of giant squid, which are hunted in the ocean depths
largely using the whale’s acoustic echolocation sense. Orcas (killer whales)
effectively use pack hunting techniques to hunt larger whales and other
marine mammals.
The deeper regions of the ocean are dominated by different types of
nekton. However, we know even less about their ecology due to their re-
lative inaccessibility. The disphotic or mesopelagic zone contains many
animal species that migrate vertically into surface waters at night to feed
upon the plankton there. Many of these organisms possess large, well-
developed eyes and also possess light organs containing symbiotic lumi-
nescent bacteria. The majority of the fish species in this group are colored
black and the invertebrates are largely red (red light penetrates water less
effectively than do longer wavelengths, and these animals appear dark-
colored at depth). Beneath this zone, in the bathypelagic and abyss-
alpelagic zones, there are many fewer organisms and much less diversity
than in the shallower levels. Animals in this region are typically colorless
and possess small eyes and luminescent organs. Because organisms in
these deep regions are few and far between, many species have become
specialized in order to maximize their advantages. Thus, deep-sea fish are
characterized by large teeth and remarkably hinged jaws that allow them
to consume prey much larger than might be expected from their size. Simi-
larly, since encounters with potential mates are presumably scarce, a num-
ber of unique reproductive strategies have evolved. In the anglerfish
(Ceratius), all of the large individuals are female and the comparatively
tiny males are parasitic, permanently attaching themselves to the female.
Much, however still remains to be learned of the ecology of these deep-sea
organisms.
396
Marine biomes
Benthos
The benthos of the world’s oceans consists of animals that live on the solid
substrate of the water column, the ocean floor. Scientists typically divide
benthic organisms into two categories, the epifauna, which live on the sur-
face of the bottom at the sediment-water interface and the infauna, those
organisms living within the sediments. In shallow water benthic commu-
nities, members of virtually every major animal group are represented.
Ecologists generally differentiate between soft bottom benthic communi-
ties (sand, silt, and mud, which comprise the majority of the benthic zone)
and rocky bottom communities, which are less common proportionately.
Soft bottom communities have an extensive diversity of burrowing
infauna, such as polychaete worms, and mollusks, such as clams. Rocky
bottom communities possess a larger proportion of epifauna, such as crus-
taceans and echinoderms (starfish, sea urchins, and brittle stars), living on
the surface of what is essentially a two-dimensional environment. Vertical
faces of the hard bottom environment, such as canyon walls or coral reefs,
are often home to a wide variety of animals occupying various crannies
and caves. In some parts of the world, kelp plants that are anchored to the
substrate and which extend to the water surface dominate the rocky bot-
tom substrate. In these kelp forests, large kelp plants (actually a species of
brown algae) form a forestlike canopy that plays host to a wide and com-
plex array of animals extending throughout the water column. On the
deep ocean floor, the benthos is composed of representatives of virtually
major animal group: crustaceans such as amphipods, segmented poly-
chaete worms, sea cucumbers, and brittle stars. Less common are starfish,
sea lilies, anemones, and sea fans. The fish of the deep benthos include rat
tails and a number of eel species.
Estuaries, where freshwater rivers empty into marine environments,
are typified by large, cyclic changes in temperature and salinity. Although
estuaries have played an important role in human history as the sites of
major ports, the variety and number of estuarine species tend to show less
diversity of animal species due to the difficulty in adapting to the large
swings in environmental conditions.
Animal life in the sea, like that on land, shows an astonishing variety of
forms and behaviors, the result of natural selection. The inaccessibility and
hostility of much of the world’s oceans to human exploration and observa-
tion leaves much yet to be learned about the biology of marine life. Much
remains to be achieved in order to obtain a useful body of knowledge con-
cerning life in the sea.
John G. New
397
Marine biomes
See also: Acid deposition; Eutrophication; Evolution: definition and theo-

ries; Habitats and biomes; Invasive plants; Lakes and limnology; Ocean
pollution and oil spills; Reefs; Wetlands.

Niesen, T. M. The Marine Biology Coloring Book. 2d ed. New York: Harper-
Resource, 2000.
Nybakken, J. W. Marine Biology: An Ecological Approach. 5th ed. San Fran-
cisco: Benjamin/Cummings, 2001.
Robison, B. H., and J. Connor. The Deep Sea. Monterey Bay, Calif.: Monterey
Bay Aquarium Press, 1999.
Safina, C. Song for the Blue Ocean: Encounters Along the World’s Coasts and Be-
neath the Seas. New York: Henry Holt, 1998.
398
MEDITERRANEAN SCRUB
Mediterranean scrub vegetation is dominated by fire-adapted shrubs. The biome

fringes the Mediterranean Sea, for which it is named, but is also found along west-
ern coasts of continents in areas with warm, dry summers and moist, cool winters.
R egions with mediterranean vegetation are coastal regions between 30

and 45 degrees north latitude or between 30 and 45 degrees south lati-
tude. The air circulating around high-pressure zones over adjacent oceans
guides storms away from the coast in the warm season but changes posi-
tion in concert with the tilt of the earth on its axis and brings storms onto
the coast in the cool season. As a result, the warm season is dry, and the
cool season is moist. Fire is an important component of mediterranean en-
vironments, especially after the warm, dry summer.
North America’s representative of mediterranean scrub is the chaparral
of the Pacific Coast of Southern California and northern Baja California,
Mexico. In chaparral and some other mediterranean regions, winds blow-
ing from continental high-pressure regions toward the coast help push
storm tracks offshore during the warm season. In California these winds
are called Santa Ana winds and are best known for driving chaparral fires.
Lightning started such fires before human settlement, but they are often
started by careless people today. With the cooler temperatures of autumn
and winter the continental pressure wanes and the Santa Ana winds de-
crease. At the same time, the oceanic high-pressure region shifts, and win-
ter storms track onto the coast, bringing the cool season rains.
Character and Components

Mediterranean scrub is found in small, scattered areas around the world.
The plant species that occur in this biome on one continent are unrelated to
those that occur in the same biome on other continents. As a result, medi-
terranean scrub presents a classical example of convergent evolution, the
environmentally driven development of similar characteristics in unre-
lated species. Under the influence of mediterranean climate, entire com-
munities of unrelated species become similar to one another. Many medi-
terranean areas also contain a large number of endemic plant species,
species that grow nowhere else.
Mediterranean scrub is dominated by shrubs well adapted to fire. Some
species have specialized underground structures that are undamaged by
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Mediterranean scrub
the fire and send up new growth shortly after the fire passes. Other species
have specialized, long-lived seeds that require intense heat to stimulate
germination. Still other species combine the two strategies. In communi-
ties that burn regularly, such species have a great advantage over their
competitors.
Mediterranean shrubs are not just adapted to recover after a fire; they
are actually adapted to carry the fire once it is started. These species syn-
thesize and store highly flammable chemicals in their leaves and stems.
The flammable vegetation ensures that most fires will burn large areas.
The most widespread shrub in North American chaparral is chamise
(Adenostoma fasciculatum), which sprouts from underground structures
and produces large numbers of seedlings after a fire. Various species of
manzanita (Arctostaphylos) and wild lilac (Ceanothus) are also widespread
throughout chaparral. Some species in each genus both sprout and pro-
duce large numbers of seedlings after fires. Other species in each genus de-
pend entirely on heat-stimulated seeds to reestablish their presence in a
burned area.
Mediterranean vegetation also occurs on western coasts in southern
Australia, where it is called mallee; the Cape region of South Africa (fyn-
bos); the central coast of Chile (matorral); and around the Mediterranean
Sea (maquis). In all these areas, the vegetation has the same adaptive char-
acteristics and appearance, but the species are not related to those of other
areas. Although there are differences among the regions besides the spe-
cies that occur in each, the similar physical and vegetational characteris-
tics lend a continuity that is widely recognized as the mediterranean scrub
biome.
Human Impact
As people moved into Mediterranean scrub regions, two major and related
concerns surfaced. First, the fires, which are such an important part of
scrub ecology, were destructive and dangerous, leading to fire suppres-
sion. Second, fire suppression may actually increase fire damage and may
threaten the mediterranean scrub biome’s very existence when combined
with other human activities. A comparison of the fire history in the chapar-
ral of California and that of Baja California lends credibility to the idea that
fire suppression increases fire damage. Fire suppression has long been
practiced in Southern California. In contrast, much less fire suppression
has gone on in Baja. Fewer, larger, and more destructive fires burn in
Southern California chaparral than in Baja chaparral. The simplest expla-
nation is that fire suppression allows fuel to build up, so that when a fire
starts it is essentially unstoppable, as often occurs in California chaparral.
400
Mediterranean scrub
With less fire suppression and less fuel accumulation, Baja fires burn more
frequently but are smaller and less destructive. The small fires remove the
fuel periodically, thus decreasing the danger of large, destructive fires.
There are other differences between California and Baja chaparral that
may account for the differences in the fire regimes, but the foregoing hy-
pothesis is interesting from the perspective of human impact on chaparral
as well as that of fire’s impact on humans. Population growth and its atten-
dant activities threaten the very existence of the chaparral. Humans de-
stroy chaparral to build home sites, suppress fires, and plant grass in
burned areas to stabilize the soil and to mitigate future fires. The grasses
compete with chaparral plants and retard chaparral recovery. The impact
of these and other activities on the native chaparral ecosystem is not well
understood but is almost certainly negative. Other mediterranean scrub
areas suffer similar fates. Although mediterranean scrub is still well repre-
sented in comparison to some biomes, its response to human impact
should be carefully studied and monitored, both to protect human invest-
ment in mediterranean ecosystems and to preserve the intriguing mediter-
ranean scrub and its many unique plant species.
Carl W. Hoagstrom
See also: Biomes: determinants; Biomes: types; Chaparral; Forest fires; For-
est management; Forests.

Barbour, Michael G., and William Dwight Billings, eds. North American Ter-
restrial Vegetation. 2d ed. New York: Cambridge University Press, 2000.
Dallman, Peter R. Plant Life in the World’s Mediterranean Climates. Berkeley:
University of California Press, 1998.
Vankat, John L. The Natural Vegetation of North America: An Introduction.
Melbourne, Fla.: Krieger, 1992.
401
METABOLITES
Type of ecology: Chemical ecology
Metabolites are compounds synthesized by plants for both essential functions,

such as growth and development (primary metabolites), and specific functions,
such as pollinator attraction of defense against herbivory (secondary metabolites).
M etabolites are organic compounds synthesized by organisms using

enzyme-mediated chemical reactions called metabolic pathways.
Primary metabolites have functions that are essential to growth and devel-
opment and are therefore present in all plants. In contrast, secondary metab-
olites are variously distributed in the plant kingdom, and their functions
are specific to the plants in which they are found. Secondary metabolites
are often colored, fragrant, or flavorful compounds, and they typically me-
diate the interaction of plants with other organisms. Such interactions in-
clude those of plant-pollinator, plant-pathogen, and plant-herbivore.
Primary Metabolites
Primary metabolites comprise many different types of organic compounds,
including, but not limited to, carbohydrates, lipids, proteins, and nucleic
acids. They are found universally in the plant kingdom because they are
the components or products of fundamental metabolic pathways or cycles
such as glycolysis, the Krebs cycle, and the Calvin cycle. Because of the im-
portance of these and other primary pathways in enabling a plant to syn-
thesize, assimilate, and degrade organic compounds, primary metabolites
are essential.
Examples of primary metabolites include energy-rich fuel molecules,
such as sucrose and starch, structural components such as cellulose, in-
formational molecules such as DNA (deoxyribonucleic acid) and RNA (ri-
bonucleic acid), and pigments, such as chlorophyll. In addition to having
fundamental roles in plant growth and development, some primary me-
tabolites are precursors (starting materials) for the synthesis of secondary
metabolites.
Secondary Metabolites
Secondary metabolites largely fall into three classes of compounds: alka-
loids, terpenoids, and phenolics. However, these classes of compounds
also include primary metabolites, so whether a compound is a primary
or secondary metabolite is a distinction based not only on its chemi-
402
Metabolites
cal structure but also on its function and distribution within the plant
kingdom.
Many thousands of secondary metabolites have been isolated from
plants, and many of them have powerful physiological effects in humans
and are used as medicines. It is only since the late twentieth century that
secondary metabolites have been clearly recognized as having important
functions in plants. Research has focused on the role of secondary metabo-
lites in plant defense. This is discussed below with reference to alkaloids,
though it is relevant to many types of secondary metabolites.
Alkaloids
Alkaloids are a large group of nitrogen-containing compounds, examples
of which are known to occur in approximately 20 percent of all flowering
plants. Closely related plant species often contain alkaloids of related
chemical structure. The primary metabolites from which they are derived
include amino acids such as tryptophan, tyrosine, and lysine. Alkaloid
biosynthetic pathways can be long, and many alkaloids have correspond-
ingly complex chemical structures. Alkaloids accumulate in plant organs
such as leaves or fruits and are ingested by animals that consume those
plant parts. Many alkaloids are extremely toxic, especially to mammals,
and act as potent nerve poisons, enzyme inhibitors, or membrane trans-
port inhibitors. In addition to being toxic, many alkaloids are also bitter or
otherwise bad-tasting. Therefore, the presence of alkaloids and other toxic
secondary metabolites can serve as a deterrent to animals to avoid eating
such plants.
Sometimes domesticated animals that have not previously been ex-
posed to alkaloid-containing plants do not have acquired avoidance mech-
anisms, and they become poisoned. For example, groundsel contains the
alkaloid senecionine, which has resulted in many recorded cases of live-
stock fatalities due to liver failure. More frequently, over time, natural se-
lection has resulted in animals developing biochemical mechanisms or be-
havioral traits that lead to avoidance of alkaloid-containing plants.
In other, more unusual cases, animals may evolve a mechanism for se-
questering (storing) or breaking down a potentially toxic compound, thus
“disarming” the plant. For instance, caterpillars of the cinnabar moth can
devour groundsel plants and sequester senecionine without suffering any
ill effects. Moreover, the caterpillars thereby acquire their own weapon
against predators: the plant-derived alkaloid stored within their bodies.
Over time, plants acquire new capabilities to synthesize additional defense
compounds to combat animals that have developed “resistance” to the
original chemicals. This type of an “arms race” is a form of coevolution and
403
Metabolites
may help to account for the incredible abundance of secondary metabolites

in flowering plants.
Medicinal Alkaloids
Many potentially toxic plant-derived alkaloids have medicinal properties,
as long as they are administered in carefully regulated doses. Alkaloids
with important medicinal uses include morphine and codeine from the
opium poppy and cocaine from the coca plant. These alkaloids act on the
nervous system and are used as painkillers. Atropine, from the deadly
nightshade plant, also acts on the nervous system and is used in anesthesia
and ophthalmology. Vincristine and vinblastine from the periwinkle plant
are inhibitors of cell division and are used to treat cancers of the blood and
lymphatic systems. Quinine from the bark of the cinchona tree is toxic to
the Plasmodium parasite, which causes malaria, and has long been used in
tropical and subtropical regions of the world. Other alkaloids are used as
stimulants, including caffeine, present in coffee, tea, and cola plants (and
the drinks derived from these plants); and nicotine, which is present in to-
bacco. Nicotine preparations are, paradoxically, also used as an aid in
smoking cessation. Nicotine is also a very potent insecticide. For many
years ground-up tobacco leaves were used for insect control, but this prac-
tice was superseded by the use of special formulations of nicotine. More re-
cently the use of nicotine as an insecticide has been discouraged because of
its toxicity to humans.
Terpenoids
Terpenoids are derived from acetyl coenzyme A or from intermediates in
glycolysis. They are classified by the number of five-carbon isoprenoid
units they contain. Monoterpenes (containing two C5-units) are exempli-
fied by the aromatic oils (such as menthol) contained in the leaves of mem-
bers of the mint family. In addition to giving these plants their characteris-
tic taste and fragrance, these aromatic oils have insect-repellent qualities.
The pyrethroids, which are monoterpene esters from the flowers of chry-
santhemum and related species, are used commercially as insecticides.
They fatally affect the nervous systems of insects while being biodegrad-
able and nontoxic to mammals, including humans.
Diterpenes are formed from four C5-units. Paclitaxel (commonly known
by the name Taxol), a diterpene found in bark of the Pacific yew tree,
is a potent inhibitor of cell division in animals. At the end of the twenti-
eth century, paclitaxel was developed as a powerful new chemothera-
peutic treatment for people with solid tumors, such as ovarian cancer pa-
tients.
404
Metabolites
Triterpenoids (formed from six C5-units) comprise the plant steroids,

some of which act as plant hormones. These also can protect plants from
insect attack, though their mode of action is quite different from that of the
pyrethroids. For example, the phytoecdysones are a group of plant sterols
that resemble insect molting hormones. When ingested in excess, phyto-
ecdysones can disrupt the normal molting cycle with often lethal conse-
quences to the insect.
Tetraterpenoids (eight C5-units) include important pigments such as
beta-carotene, which is a precursor of vitamin A, and lycopene, which
gives tomatoes their red color. Rather than functioning in plant defense,
the colored pigments that accumulate in ripening fruits can serve as attrac-
tants to animals, which actually aid the plant in seed dispersal.
The polyterpenes are polymers that may contain several thousand
isoprenoid units. Rubber, a polyterpene in the latex of rubber trees that
probably aids in wound healing in the plant, is also very important for the
manufacture of tires and other products.
Phenolic Compounds
Phenolic compounds are defined by the presence of one or more aromatic
rings bearing a hydroxyl functional group. Many are synthesized from the
amino acid phenylalanine. Simple phenolic compounds, such as salicylic
acid, can be important in defense against fungal pathogens. Salicylic acid
concentration increases in the leaves of certain plants in response to fungal
attack and enables the plant to mount a complex defense response. Inter-
estingly, aspirin, a derivative of salicylic acid, is routinely used in humans
to reduce inflammation, pain, and fever. Other phenolic compounds, called
isoflavones, are synthesized rapidly in plants of the legume family when
they are attacked by bacterial or fungal pathogens, and they have strong
antimicrobial activity.
Lignin, a complex phenolic macromolecule, is laid down in plant sec-
ondary cell walls and is the main component of wood. It is a very impor-
tant structural molecule in all woody plants, allowing them to achieve
height, girth, and longevity. Lignin is also valuable for plant defense: Plant
parts containing cells with lignified walls are much less palatable to insects
and other animals than are nonwoody plants and are much less easily di-
gested by fungal enzymes than plant parts that contain only cells with pri-
mary cellulose walls.
Other phenolics function as attractants. Anthocyanins and antho-
cyanidins are phenolic pigments that impart pink and purple colors to
flowers and fruits. This pigmentation attracts insects and other animals
that move between individual plants and accomplish pollination and fruit
405
Metabolites
dispersal. Often the plant pigment and the pollinator’s visual systems are
well matched: Plants with red flowers attract birds and mammals because
these animals possess the correct photoreceptors to see red pigments.
Valerie M. Sponsel
See also: Allelopathy; Defense mechanisms; Genetically modified foods;

Pheromones; Poisonous animals; Poisonous plants.

Levetin, Estelle, and Karen McMahon. Plants and Society. Boston: WCB/
McGraw-Hill, 1999.
Moore, Randy, et al. Botany. 2d ed. Boston: WCB/McGraw-Hill, 1998.
406
MIGRATION
Migration is an important ecological process that results in the redistribution of

animal populations from one habitat to another. Adaptive habitat changes are fun-
damentally important aspects of the life histories of many species.
M igration is a general term employed by ecologists and ethologists to

describe the nearly simultaneous movement of many individuals or
entire populations of animals to or between different habitats. As defined,
migrations do not include local excursions made by individuals or small
groups of animals in search of food, to mark territorial boundaries, or to ex-
plore surrounding environments.
Nomads are migrants whose populations follow those of their primary
food sources. Such animals (the American bison, for example) do not have
fixed home ranges and wander in search of suitable forage. Some scientists
view nomadic movements as a form of extended foraging behavior rather
than as a special case of migration. In either context, the important point is
that populations change habitats in response to changing conditions.
In contrast to migrations made by populations and excursions made by
individuals, the spreading or movement of animals away from others is
known as dispersal. Examples of dispersal include the drift of plankton in
currents and the departure of subadult animals from the home range of
their parents. In numerous species (sea turtles, rattlesnakes, and salmon,
for example), dispersed members of a population may return to the place
of origin after a variable interval of time.
Navigation
Among the animals known to navigate are birds (the best-studied group),
lobsters, bees, tortoises, bats, marine and terrestrial mammals, fish, brittle
starfish, newts, toads, and insects. Some migratory species can orient
themselves—that is, they know where they are in time and space. Many
birds and mammals, for example, have an inherent sense of the direction,
distance, and location of distant habitats. Orientation and travel along un-
familiar routes from one place or habitat to another is called navigation.
Navigators use environmental and sensory information to reach distant
geographical locations, and many of them do so with a remarkably accu-
rate sense of timing. Homing pigeons are perhaps the best-studied animal
navigators. These birds are able not only to discover where they are when
407
Migration
released but also to return to their home loft from distant geographical lo-
cations.
Much has been learned about how animals successfully navigate over
long distances from the pioneering studies of Archie Carr. Carr proposed
that green sea turtles successfully find their widely separated nesting and
feeding beaches by means of an inherent clock sense, map sense, and com-
pass sense. His investigations and those of many others continue to stimu-
late great interest in the physiology and ecology of navigating species and
in the environmental cues to which they respond. Sensory biologists, bio-
physicists, and engineers have incorporated knowledge of how animals
detect and use environmental information to develop new and more accu-
rate navigational systems for human use.
Animals use a variety of cues to locate their positions and appropriate
travel paths. Most species have been found to use more than one type of in-
formation (sequentially, alternatively, or simultaneously) to navigate. In-
cluded among the orientation guideposts that one or more of these groups
may use are the positions of the sun and stars, magnetic fields, ultraviolet
light, tidal fluctuations caused by the changing positions of the moon and
sun, atmospheric pressure variations, infrasounds (very low frequency
sounds), polarized light (on overcast days), environmental odors, shore-
line configurations, water currents, and visual landmarks. Celestial cues
also require a time sense, or an internal clock, to compensate for move-
ments of the animal relative to changing positions of celestial objects in the
sky. In addition to an absolute dependence on environmental cues, young
or inexperienced members of some species may learn navigational routes
from experienced individuals, such as their parents, or other experienced
individuals in the population. Visual mapping remembered from explor-
atory excursions may also play a role in enhancing the navigational abili-
ties of some birds, fish, mammals, and other animals.
Benefits of Migration
The different categories of animal movements, however, are perhaps not as
important as the reasons animals migrate and the important biological
consequences of the phenomenon. As a general principle, migrations are
adaptive behavioral responses to changes in ecological conditions. Popula-
tions benefit in some way by regularly or episodically moving from one
habitat to another.
An example of the adaptive value of migratory behavior is illustrated
by movement of a population from a habitat where food, water, space,
nesting materials, or other resources have become scarce (often a seasonal
phenomenon) to an area where resources are more abundant. Relocation to
408
Migration
a new habitat (or to the same type of habitat in a different geographical

area) may reduce intraspecific or interspecific competition, may reduce
death rates, and may increase overall fitness in the population. These bene-
fits may result in an increase in reproduction in the population. Reproduc-
tive success, then, is the significant benefit and the only biological criterion
used to evaluate population fitness.
Programmed Movements
While many factors are believed to initiate migratory events, most fall into
one of two general categories. The first and largest category may be called
programmed movements. Such migrations usually occur at predictable in-
tervals and are important characteristics of a species’ lifestyle or life cycle.
Programmed migrations are not, in general, density-dependent. Move-
ments are not caused by overcrowding or other stresses resulting from an
excessive number of individuals in the population.
The lifestyle of a majority of drifting animals whose entire lives are
spent in the water column, for example, includes a vertical migration from
deep water during the day to surface waters at night. Thus, plankton ex-
hibit a circadian rhythm (activity occurring during twenty-four-hour inter-
vals) in their movements. An abundance of food at or near the surface and
escape from deep-water predators are among the possible reasons for
these migrations. Daily vertical movements of plankton are probably initi-
ated by changes in light intensity at depth, and the animals follow light
levels as they move toward the surface with the sinking sun. It is interest-
ing to note that zooplankters living in polar waters during the winter-long
night do not migrate.
Monarch butterflies and many large, vertebrate animals, such as her-
ring, albatross, wildebeests, and temperate-latitude bats, migrate from one
foraging area to another, or from breeding to foraging habitats, on a sea-
sonal or annual basis. Annual migrations usually coincide with seasonal
variation. Changes in day length, temperature, or the abundance of pre-
ferred food items associated with seasonal change may stimulate mass
movements directly, or indirectly, through hormonal or other physiologi-
cal changes that are correlated with seasonal environmental change. The
onset of migration in many vertebrates is evidenced by an increase in rest-
lessness that seems, in human terms, to be anticipatory.
In addition to their daily vertical migrations (lifestyle movements), the
life cycles of marine zooplankton involve migrations, and it is convenient
to use them as examples. As discussed, most adult animal plankters are
found at depth during the day and near the surface at night. In contrast,
zooplankton eggs and larvae remain in surface waters both day and night.
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Migration
Bird migrations are among the most noticeable of animal migrations, with huge
flocks passing through the skies on their way to a seasonal home that can be thou-
sands of miles distant from their original location. (Corbis)
As the young stages grow, molt, and change their shapes and food sources,
they begin to migrate vertically. The extent of vertical migrations gradually
increases throughout the developmental period, and as adults, these ani-
mals assume the migratory patterns of their parents. Patterns of movement
that change during growth and development are examples of ontogenetic,
or life-cycle, migrations.
Episodic Movements
The second large category of migratory behavior includes episodic, density-
dependent population movement. Such migrations are often associated
with, or caused by, adverse environmental changes (effect) that may be
caused by overlarge populations (cause). Local resources are adequate to
support a limited number of individuals (called the carrying capacity of
the environment), but once that number has been exceeded, the population
must either move or perish. Unfortunately, migration to escape unfavor-
able conditions may be unsuccessful, as another suitable habitat may not
be encountered. Migrations caused by overpopulation or environmental
degradation are common. Pollution and habitat destruction by human-
kind’s activities are increasingly the cause of degraded environments, and
410
Migration
in such cases, it is reasonable to conclude that humans have reduced the

carrying capacity of many animal habitats. Familiar examples of density-
dependent migrations are those of lemmings, locusts, and humans.
Ecological Import
Environmental or physiological factors that initiate migrations may be of
interest to sensory biologists and physiological ecologists; knowledge of
variation in population distributions is important to biogeographers and
wildlife biologists; and migrations in predator-prey relationships, compe-
tition, pollution, and life-history strategies are important aspects of classi-
cal ecological studies.
In addition to the specific aspect of migration being studied, the partic-
ular group of animals under investigation (moths, eels, elephants, snails)
requires that different methods be used. Some of the approaches used in
migration-related research illustrate how information and answers are ob-
tained by scientists.
Arctic terns migrate from their breeding grounds in the Arctic to the
Antarctic pack ice each year. The knowledge that these birds make a
twenty-thousand-mile annual round-trip comes from the simplest and
most practical method: direct observation of the birds (or their absence) at
either end of the trip. Direct observation by ornithologists of the birds in
flight can establish what route they take and whether they pause to rest or
feed en route. Many birds have also been tracked using radar or by obser-
vations of their silhouettes passing in front of the moon at night. Birds are
often banded (a loose ring containing coded information is placed on one
leg) to determine the frequency of migration and how many round-trips an
average individual makes during its lifetime. From this information, esti-
mates of longevity, survivorship rates, and nesting or feeding site prefer-
ences can be made.
Factors that initiate migratory behavior in terns and in other birds
can often be determined by ecologists able to relate environmental condi-
tions (changes in temperature, day length, and the like) to the timing of mi-
grations. Physiological ecologists study hormonal or other physiological
changes that co-occur with environmental changes. Elevated testosterone
levels, for example, may signal the onset of migratory behavior.
How Arctic terns orient and navigate along their migratory routes is
usually studied by means of laboratory-conducted behavioral experi-
ments. Birds are exposed to various combinations of stimuli (magnetic
fields, planetarium-like celestial fields, light levels), and their orientation,
activity levels, and physiological states are measured. Experiments involv-
ing surgical or chemical manipulation of known sensory systems are
411
Migration
sometimes conducted to compare behavioral reactions to experimental

stimuli. In such experiments, the birds (or other test animals) are rarely
harmed.
Tags of several types are used to study migrations in a wide variety of
animals, including birds, bees, starfish, reptiles, mammals, fish, snails, and
many others. Tags may be transmitting collars (located by direction-finding
radio receivers); plastic or metal devices attached to ears, fins, or flippers;
or even numbers, painted on the hard exoskeletons of bees and other in-
sects. Additional types of tagging (or identifying) include radioactive im-
plants and microchips that can be read by computerized digitizers; the use
of brands and tattoos; and, of great interest, the use of biological tags. Para-
sites known to occur in only one population of migrants (nematode para-
sites of herring, for example) provide an interesting illustration of how the
distribution of one species can be used to provide information about an-
other.
The Importance of Migration

The causes, frequency, and extent of animal migration are so diverse that
several definitions for the phenomenon have been proposed. None of
these has been accepted by all scientists who study animal movements,
however, and it is sometimes difficult to interpret what is meant when the
term “migration” is used. Most researchers have adopted a broad compro-
mise to include all but trivial population movements that involve some de-
gree of habitat change.
It is important to recognize that few populations of animals are static;
even sessile animals (such as oysters and barnacles) undergo developmen-
tal habitat changes, which are referred to as ontogenetic migrations. Aside
from certain tropical and evergreen forest areas where migrations are rela-
tively uncommon, a significant number of both aquatic and terrestrial spe-
cies move from one habitat to another at some time during their lives. In
the face of environmental change, including natural events such as sea-
sonal variation and changes caused by resource limitations and environ-
mental degradation, animals must either move, perish, or escape by means
of drastic population reduction or by becoming inactive until conditions
become more favorable (hibernation, arrested development and dormancy,
and diapause in insects are examples of behavioral-ecological inactivity).
Migration is the most common behavioral reaction to unfavorable environ-
mental change exhibited by animals.
One cannot understand the biology of migrators until their distribution
and habitats throughout life are known. The patterns of animal move-
ments are fascinating, and it is useful to summarize some of the major dif-
412
Migration
ferences between them. First, many species travel repeatedly during their
lives between two habitats, on a daily basis (as plankton and chimney
swifts do) or on an annual basis (as frogs and elks do). Second, some spe-
cies migrate from one habitat (usually suitable for young stages) to another
(usually the adult habitat) only once during their lives (for example, sal-
mon, eels, damselflies, and most zooplankton, which live on the bottom as
adults). Third, some species (many butterflies, for example) are born and
mature in one geographical area (England, for example), migrate as adults
to a distant geographical area (Spain, for example), and produce offspring
that mature in the second area. These migrations take place between gener-
ations. In a fourth pattern, one may include the seasonal swarming of so-
cial arthropods such as termites, fire ants, and bees. A fifth but ill-defined
pattern is discernible, exemplified by locust “plagues,” irruptive emigra-
tion in lemmings and certain other rodents, and some mass migrations by
humankind, as caused by war, famine, fear, politics, or disease. These are
episodic and often, if not primarily, caused by severe population stress or
catastrophic environmental change.
Sneed B. Collard
See also: Altruism; Communication; Defense mechanisms; Displays; Ethol-

ogy; Habituation and sensitization; Herbivores; Hierarchies; Insect societ-
ies; Isolating mechanisms; Mammalian social systems; Mimicry; Omni-
vores; Pheromones; Poisonous animals; Predation; Reproductive strategies;
Territoriality and aggression.

Able, Kenneth P., ed. Gatherings of Angels: Migrating Birds and Their Ecology.
Ithaca, N.Y.: Comstock, 1999.
Aidley, David, ed. Animal Migration. New York: Cambridge University
Press, 1981.
Begon, Michael, John Harper, and Colin Townsend. Ecology: Individuals,
Populations, and Communities. 3d ed. Sunderland, Mass.: Sinauer Associ-
ates, 1996.
Dingle, Hugh. Migration: The Biology of Life on the Move. New York: Oxford
Eisner, Thomas, and Edward Wilson, eds. Animal Behavior: Readings from
“Scientific American.” San Francisco: W. H. Freeman, 1975.
Newberry, Andrew, ed. Life in the Sea: Readings from “Scientific American.”
San Francisco: W. H. Freeman, 1982.
Pyle, Robert Michael. Chasing Monarchs: Migrating with Butterflies of Pas-
sage. Boston: Houghton Mifflin, 1999.
413
Migration
Rankin, Mary, ed. Migration: Mechanisms and Adaptive Significance. Port

Aransas, Tex.: Marine Science Institute, University of Texas at Austin,
1985.
Reader’s Digest Association. The Wildlife Year. Pleasantville, N.Y.: Author,
1993.
Schone, Hermann. Spatial Orientation: The Spatial Control of Behavior in Ani-
mals and Man. Translated by Camilla Strausfeld. Princeton, N.J.: Prince-
414
MIMICRY
Types of ecology: Behavioral ecology; Physiological ecology
Mimicry is the process whereby one organism resembles another and, because of
this resemblance, obtains an evolutionary advantage.
T he broadest description of mimicry is when one organism, called the

operator or dupe, cannot distinguish a second organism, referred to as
the mimic, from a third organism or a part of the environment, called the
model. There are many different types of mimicry. Some mimics look like
another organism; some smell like another organism; some may even feel
like another organism. There are also many ways that mimicking another
organism could be helpful. Mimicry may help to hide an organism in plain
sight or protect a harmless organism from predation when it mimics a
harmful organism. It can even help predators sneak up on prey species
when the predator mimics a harmless organism.
Camouflage vs. Mimicry

In the case of hiding in plain sight, the line between camouflage and mim-
icry is not sharply defined. Spots or stripes that help an organism blend
with the surroundings is classified as camouflage, because those patterns
allow the organism to remain hidden in many areas that have mixtures of
sunlight and shadow, and the organism does not look like any particular
model. As an organism’s appearance begins to mimic another organism
more and more closely, rather than displaying just a general pattern, it
moves toward mimicry. As in all other areas of biology, there are argu-
ments about where camouflage ends and mimicry begins. The stripes of a
tiger and the spots on a fawn are certainly camouflage. The appearance of a
stick insect is more ambiguous. Its body is very thin and elongated and is
colored in shades of brown and gray. Is this mimicry of a twig or just very
good camouflage? Many biologists disagree. The shapes and colors of
many tropical insects, especially mantises, also fall into this gray area of ei-
ther extremely good camouflage or simple mimicry.
Coloration: Batesian and Müllerian Mimicry

In contrast to camouflage, which hides its bearers, many species of danger-
ous or unpalatable animals are brightly colored. This type of color pattern,
which stands out against the background, is called warning coloration.
Some examples are the black and white stripes of the skunk, the yellow
415
Mimicry
and black stripes of bees and wasps, red, black, and yellow stripes of the
coral snake, and the bright orange of the monarch butterfly. Several species
of harmless insects have the same yellow and black pattern that is seen on
wasps. In addition to mimicking the coloration of the more dangerous in-
sects, some harmless flies even mimic the wasps’ flying patterns or their
buzzing sound. In each case, animals that have been stung by wasps or
bees avoid both the stinging insects and their mimics. This mimicry of
warning coloration is called Batesian mimicry. Batesian mimicry is also
seen in the mimicry of the bright red color of the unpalatable red eft stage
of newts by palatable salamanders.
Sometimes two or more dangerous or unpalatable organisms look very
much alike. In this case, both are acting as models and as mimics. This
mimicry is called Müllerian mimicry. Müllerian mimicry is seen in mon-
arch and viceroy butterflies. Both butterflies have in their bodies many of
the chemicals found in the plants they ate as larvae. These include many
unpalatable chemicals and even toxic chemicals that cause birds to vomit.
If a bird eats either a monarch or a viceroy that has these chemicals, the bird
usually remembers and avoids preying on either species again—a classic
Monarch butterflies and

viceroy butterflies both have in
their bodies unpalatable and
even toxic chemicals that cause
birds to vomit and avoid
preying on them later—a
classic example of Müllerian
mimicry, in which two or more
dangerous or unpalatable
organisms look very much
alike. It is believed that
evolutionary processes select
for such mimicry because it
simplifies the process of
recognition and avoidance on
the part of predators, thereby
increasing the fitness of the
similar-looking prey.
(PhotoDisc)
416
Mimicry
Müllerian mimicry. Interestingly, not all monarchs or viceroys are unpalat-

able. It depends on the types and concentrations of chemicals in the partic-
ular plants on which they fed as larvae. Birds that have eaten the palatable
monarchs or viceroys do not reject either monarchs or viceroys when of-
fered them as food, but birds that have eaten an unpalatable monarch or an
unpalatable viceroy avoid both palatable and unpalatable members of
both species. This represents both Batesian and Müllerian mimicry at
work.
Aggressive Mimicry
Mimicry by predators is called aggressive mimicry. The reef fish, called the
sea swallow, is a cleaner fish, and larger fish enter the sea swallow’s terri-
tory to be cleaned of parasites. The saber-toothed blenny mimics the
cleaner in both appearance and precleaning behavior, but when fish come
to be cleaned, the blenny instead bites off a piece of their flesh to eat.
Anglerfish have small extensions on their heads that resemble worms.
They mimic worms to lure their prey close enough to be eaten. The alliga-
tor snapping turtle’s tongue and the tips of the tails of moccasins, copper-
heads, and other pit vipers are also wormlike and are used as lures. Certain
predatory female fireflies respond to the light flashes of males of a different
species with the appropriate response of the female of that species. This
lures the male closer, and when the unsuspecting male is close enough to
mate, the female devours him. This mimicry is quite complex, because the
predatory females are able to mimic the response signals of several differ-
ent species.
Octopuses
There are many other instances of mimicry, but the world champion mim-
ics may be octopuses. As predators, these animals show unbelievable ag-
gressive mimicry of other reef organisms. Octopuses can take on the color,
shape, and even texture of corals, algae, and other colonial reef dwellers.
As a prey species, the octopus can use the same type of mimicry for camou-
flage, but can also be a Batesian mimic, taking on the color and shape of
many of the reef’s venomous denizens.
Since in each case, being a mimic helped the organism in some way, it is
not hard to understand how mimicry may have evolved. In a population in
which some organisms were protected by being mimics, the protected
mimics were most likely to mate and leave their genes for the next genera-
tion while the unprotected organisms were less likely to breed.
Richard W. Cheney, Jr.
417
Mimicry
See also: Altruism; Camouflage; Communication; Defense mechanisms;

Displays; Mammalian social systems; Pheromones; Poisonous animals;
Predation; Reproductive strategies; Territoriality and aggression.

Brower, Lincoln P., ed. Mimicry and the Evolutionary Process. Chicago: Uni-
Ferrari, Marco. Colors for Survival: Mimicry and Camouflage in Nature. Char-
lottesville, Va.: Thomasson-Grant, 1993.
Owen, D. Camouflage and Mimicry. Chicago: University of Chicago Press,
1982.
Salvato, M. “Most Spectacular Batesian Mimicry.” In University of Florida
Book of Insect Records, edited by T. J. Walker. Gainesville: University of
Florida Department of Entomology and Nematology, 1999.
Wickler, Wolfgang. Mimicry in Plants and Animals. New York: McGraw-
Hill, 1968.
418
MOUNTAIN ECOSYSTEMS
Mountains cover one fifth of the earth’s terrestrial surface, and they are one of the
most extreme environments in the global ecosystem. Mountains are globally sig-
nificant landforms that function as storehouses for irreplaceable resources such as
clean air and water, biological and cultural diversity, as well as timber and mineral
resources.
M ountains are the most conspicuous landforms on earth. They are

found on every continent and have been defined simply as elevated
landforms of high local relief, with much of the surface in steep slopes, dis-
playing distinct variations in climate and vegetation zones from the base to
the summit. The earth’s mountain ranges have been created by the colli-
sion of tectonic plates. Associated with many of these mountain ranges are
volcanoes. If the solidified magma of a volcano builds up, it can become a
mountain; likewise, if the collision involves two oceanic plates, a string of
volcanic mountains, called an island arc, can form on the ocean floor.
Mountain Habitat
Mountains are globally significant reservoirs of biodiversity. They contain
rich assemblages of species and ecosystems. Because of the rapid changes
in altitude and temperature along a mountain slope, multiple ecological
zones are stacked upon one another, sometimes ranging from dense tropi-
cal jungles to glacial ice within a few kilometers. Many plant and animal
species are found only on mountains, having evolved over centuries of iso-
lation to inhabit these specialized environments. Mountains can also func-
tion as biological corridors, connecting isolated habitats or protected areas
and allowing species to migrate between them. These extraordinary eco-
logical conditions, coupled with many bio-climatic zones, have resulted in
a high number of ecological niches available for habitation in mountain
ecosystems.
Biodiversity
Because of the great diversity in habitats within mountainous regions,
with each region showing a different combination of environmental fac-
tors, total mountain fauna is relatively rich and the variety of small com-
munities very great, in spite of the general severity of the mountain envi-
ronment as a whole. Likewise, this diversity has resulted in a wide range of
419
Mountain ecosystems
Mountains are perhaps the most significant reservoirs of biodiversity. Because of

the rapid changes in altitude and temperature along a mountain slope, multiple
ecological zones are stacked upon one another—sometimes ranging from dense
tropical jungles to glacial ice—and therefore contain rich assemblages of species
and ecosystems. (PhotoDisc)
endemic species that have evolved over centuries of isolation from other
genetic material. Rocky Mountain National Park typifies this diversity as a
home to some 900 species of plants, 250 species of birds, and 60 species of
mammals. Some are easily seen and others are elusive, but all are part of
the ecosystem in the park. On a global scale, the diversity of mountain
fauna extends to many species of ungulates, including elk, bighorn sheep,
moose, and deer. Also present in mountain communities are many species
of rodents. Rodent species may include beaver, marmots, squirrels, and
chipmunks. Other mammalian animal life include bear, canids, such as
coyote and wolf, and many species of felids, such as mountain lions and
bobcats. Mountain avian fauna comprise many families of hummingbirds,
bluebirds, hawks, falcons, eagles, and many more.
Ecological Threats
Mountains are threatened in a variety of ways, but without question, hu-
man settlement and activities such as camping, hiking, and other recre-
ational activities constitute the biggest threats to the mountain ecosystem.
420
Mountain ecosystems
Hikers and motorized offroad vehicles, for example, create tracks in the
soil that form erosion gullies and trample vegetation that has taken many
years to grow. Commercial harvesting of trees in the lower forest zones of
mountains is having an increasingly detrimental effect on biodiversity.
Global warming is another threat to mountain ecosystems. Snowlines are
receding, and continued melting of glaciers and polar ice caps could even-
tually lead to drying of major river systems which feed from them.
In an attempt to restore or conserve mountain ecosystems, many coun-
tries have replanted indigenous trees with fast-growing coniferous trees,
in an ill-fated effort to supply a growing human population with wood
products. These hybrid forests are not nearly as beautiful as the native for-
ests, but more to the point, they do not offer environments conducive to the
ecosystems that the native species supported. This loss of habitat creates a
loss of wildlife, which then becomes threatened and eventually endan-
gered because of the decline of native vegetation.
Jason A. Hubbart
See also: Biomes: types; Forest fires; Forest management; Forests; Grazing
and overgrazing; Habitats and biomes; Lakes and limnology; Old-growth
forests; Rain forests; Rain forests and the atmosphere; Restoration ecology;
Savannas and deciduous tropical forests; Sustainable development; Taiga;
Tundra and high-altitude biomes; Wildlife management.

Denniston, D. High Priorities: Conserving Mountain Ecosystems and Cultures.
Worldwatch Paper 123. Washington, D.C.: Worldwatch Institute, 1995.
Messerli, B., and J. D. Ives. Mountains of the World: A Global Priority. New
York: Parthenon, 1997.
Price, L. Mountains and Man. Berkeley: University of California Press, 1981.
Sauvain, P. Geography Detective: Mountains. Minneapolis: Carolrhoda Books,
1996.
Stronach, N. Mountains. Minneapolis: Lerner, 1995.
421
MULTIPLE-USE APPROACH
Types of ecology: Agricultural ecology; Restoration and conservation
ecology
The multiple-use approach is a concept of resource use in which land supports sev-
eral concurrent managed uses rather than single uses over time and space.
T he multiple-use approach is a management practice that is teamed

with the concept of sustained yield. Multiple use began as a working
policy, generally associated with forestry, and was enacted as law in 1960.
As a concept of land-use management, it has most often been applied to
the use of forestlands. Historically, multiple use has been linked with an-
other concept, that of sustained yield.
Historical Background
The history of the intertwined multiple-use and sustained-yield ap-
proaches to land management in the United States dates from the late
1800’s. Prior to that time, forestlands were used for timber production,
rangeland for grazing, and parklands for recreation. Little attention was
given to the interrelated aspects of land use. By the late 1800’s, however,
some resource managers began to see land as a resource to be managed in a
more complex, integrated fashion that would lead to multiple use. This
awakening grew out of the need for conservation and sustained yield, es-
pecially in the forest sector of the resource economy.
Sustained Yield
Since the earliest European settlement of North America, forest resources
had been seen both as a nearly inexhaustible source of timber and as an im-
pediment to be cleared to make way for agriculture. This policy of removal
led to serious concern by the late 1800’s about the future of American forests.
By 1891 power had been granted to U.S. president Benjamin Harrison to set
aside protected forest areas. Both he and President Grover Cleveland took
action to establish forest reserves. To direct the management of these re-
serves, Gifford Pinchot was appointed chief forester. Pinchot was trained in
European methods of forestry and managed resources, as noted by Stewart
Udall in The Quiet Crisis (1963), “on a sustained yield basis.” The sustained-
yield basis for forest management was thus established. Essentially, the
sustained-yield philosophy holds that the amount of timber harvested
should not exceed the ultimate timber growth during the same period.
422
Multiple-use approach
Multiple Use
Properly managed, forestlands can meet needs for timber on an ongoing,
renewable basis. However, land in forest cover is more than a source of
timber. Watersheds in such areas can be protected from excessive runoff
and sedimentation through appropriate management. Forest areas are also
wildlife habitat and potential areas of outdoor recreation. The combination
of forest management for renewable resource production and complex, in-
terrelated land uses provided the basis for the development of multiple-
use sustained-yield as a long-term forest management strategy.
Multiple Use Joins Sustained Yield

The merging of these two concepts took shape over a period of many years,
beginning in the early twentieth century. The establishment of national for-
ests by Presidents Harrison and Cleveland provided a base for their ex-
pansion under President Theodore Roosevelt in the early 1900’s. With
the active management of Pinchot and the enthusiastic support of Roose-
velt, the national forests began to be managed on a long-term, multiple-use
sustained-yield basis. The desirability of this approach eventually led to
its formalization by law: On June 12, 1960, Congress passed the Multiple
Use-Sustained Yield Act. To some, this act was the legal embodiment of
practices already in force. However, the act provides a clear statement
of congressional policy and relates it to the original act of 1897 that had es-
tablished the national forests.
The 1960 act specifies that “the national forests are established and shall
be administered for outdoor recreation, range, timber, watershed, and
wildlife and fish purposes.” Section 2 of the act states that the “Secretary of
Agriculture is authorized and directed to develop and administer the re-
newable resources of the national forests for multiple use and sustained
yield of the several products and services obtained therefrom.” The act
gives no specifics, providing a great deal of freedom in choosing ways to
meet its provisions. It also refrains from providing guidelines for manage-
ment. In practice, the achievement of a high level of land management un-
der the act has called for advocating a conservation ethic, soliciting citizen
participation, providing technical and financial assistance to public and
private forest owners, developing international exchanges on these man-
agement principles, and extending management knowledge.
Jerry E. Green
See also: Biopesticides; Conservation biology; Erosion and erosion control;

Genetically modified foods; Grazing and overgrazing; Integrated pest
management; Old-growth forests; Rangeland; Reforestation; Restoration
423
Multiple-use approach
ecology; Slash-and-burn agriculture; Soil; Soil contamination; Sustainable

development.

Cutter, Susan, Hilary Renwick, and William Renwick. Exploitation, Conser-
vation, Preservation: A Geographical Perspective on Natural Resource Use.
3d ed. New York: J. Wiley & Sons, 1999.
Hewett, Charles E., and Thomas E. Hamilton, eds. Forests in Demand: Con-
flicts and Solutions. Boston: Auburn House, 1982.
Lovett, Francis. National Parks: Rights and the Common Good. Lanham, Md.:
Rowman & Littlefield, 1998.
Sedjo, Roger A., ed. A Vision for the U.S. Forest Service: Goals for Its Next Cen-
tury. Washington, D.C.: Resources for the Future, 2000.
Udall, Stewart. The Quiet Crisis. 1963. Reprint. Salt Lake City, Utah: Gibbs
Smith, 1998.
424
MYCORRHIZAE
Mycorrhizae are mutualistic, symbiotic relationships between plant roots or other

underground organs and fungi. They are among the most abundant symbioses in
the world.
M ycorrhizal associations (from the Greek mukes, meaning “fungus,”

and rhiza, meaning “root”) have been described in virtually all eco-
nomically important plant groups. Investigators in Europe detected fungal
associations in most European species of flowering plants, all gymno-
sperms, ferns, and some bryophytes, especially the liverworts. Similar pat-
terns are predicted in other ecosystems. Continuing studies of ecosystems,
from boreal forests to temperate grasslands to tropical rain forests and
agroecosystems, also suggest that most plant groups are intimately linked
to one or more species of fungus.
It is theorized that most of the plants in stable habitats where competi-
tion for resources is common probably have some form of mycorrhizal as-
sociation. Species from all the major taxonomic groups of fungi, includ-
ing the Ascomycotina, Basidiomycotina, Deuteromycotina, and Zygomycotina,
have been found as partners with plants in mycorrhizae.
Considering the prevalence of mycorrhizae in the world today, bota-
nists theorize that mycorrhizae probably arose early in the development of
land plants. Some suggest that mycorrhizae may have been an important
factor in the colonization of land. The fungal partner (or mycobiont) in a
mycorrhizal relationship benefits by gaining a source of carbon. Often
these mycobionts are poor competitors in the soil environment. Some
mycobionts have apparently coevolved to the point that they can no longer
live independently of a plant host.
The plant partner in the mycorrhizal relationship benefits from im-
proved nutrient absorption. This may occur in different ways; for example,
the mycobiont may directly transfer nutrients to the root. Infected roots ex-
perience more branching, thus increasing the volume of soil that the plant
can penetrate and exploit. Evidence also suggests that mycorrhizal roots
may live longer than roots without these associations. Comparison of the
growth of plants without mycorrhizae to those with fungal partners sug-
gests that mycorrhizae enhance overall plant growth.
425
Mycorrhizae
Endomycorrhizae
Mycorrhizae may be classified into two broad groups: endomycorrhizae
and ectomycorrhizae. Endomycorrhizae enter the cells of the root cortex.
Ectomycorrhizae colonize plant roots but do not invade root cortex cells.
The most common form of endomycorrhizae are the vesicular-arbuscular
mycorrhizae. The fungi involved are zygomycetes. These mycorrhizae
have internal structures called arbuscules, which are highly branched,
thin-walled tubules inside the root cortex cells near the vascular cylinder.
It is estimated that 80 percent of all plant species may have vesicular-
arbuscular mycorrhizae. This type of mycorrhiza is especially important in
tropical trees.
There are several other subtypes of endomycorrhizae. Ericoid mycor-
rhizae, found in the family Ericaceae and closely related families, supply
the host plants with nitrogen. These are usually restricted to nutrient-poor,
highly acidic conditions, such as heath lands. Arbutoid mycorrhizae, found
in members of the Arbutoideae and related families, share some similarities
with ectomycorrhizae in that they form more developed structures called
the sheath and Hartig net (described below).
Monotropoid mycorrhizae, found in the plant family Monotropaceae, are
associated with plants that lack chlorophyll. The host plant is completely
dependent on the mycobiont, which also has connections to the roots of a
nearby tree. Thus the host, such as Monotropa, indirectly parasitizes an-
other plant by using the mycobiont as an intermediate. Orchidaceous
mycorrhizae are essential for orchid seed germination.
Ectomycorrhizae
Ectomycorrhizae are common in forest trees and shrubs in the temperate
and subarctic zones. Well-developed fungal sheaths characterize these
mycorrhizae, along with special structures called Hartig nets. Basidio-
mycetes are the usual mycobionts and often form mushrooms or truffles.
Ectomycorrhizae help protect the host plant from diseases by forming a
physical fungal barrier to infection.
The Fungal Partner

Individual filaments of a fungal body are called hyphae. The entire fungal
body is called a mycelium. Root infection may occur from fungal spores
that germinate in the soil or from fungal hyphae growing from the body of
a nearby mycorrhiza. When infection occurs, hyphae are drawn toward
certain chemical secretions from a plant root.
In ectomycorrhizae, root hairs do not develop in roots after infection oc-
curs. Infected roots have a fungal sheath, or mantle, that ranges from 20 to
426
Mycorrhizae
40 micrometers thick. Fungal hyphae penetrate the root by entering be-

tween epidermal cells. These hyphae push cells of the outer root cortex
apart and continue to grow outside the cells. This association of hyphal
cells and root cortex cells is called a Hartig net. In ectomycorrhizae, the
mycobionts never invade plant cells, nor do they penetrate the endodermis
or enter the vascular cylinder. The root tip may be ensheathed by fungi, but
the apical meristem is never invaded. Main roots experience fewer ana-
tomical changes than lateral roots after infection. Lateral roots become
thickened, may show the development of characteristic pigments, and
grow very slowly. Infected roots also show different branching patterns,
compared to those of uninfected roots.
Endomycorrhizae are highly variable in structure. Many endomycor-
rhizae do not have sheaths or Hartig nets. In all endomycorrhizae, hyphae
penetrate into root cortex cells, while portions of the mycelium remain in
contact with the soil. The hyphae that remain in the soil are important in
fungal reproduction and produce large numbers of haploid spores. Fungi do
not invade root meristems, vascular cylinders, or chloroplast-containing
cells in the plant.
Some of the host cells contain fungal extensions called vesicles that are
filled with lipids. Vesicles are specialized structures that are often thick-
walled and may serve as storage sites or possibly in reproduction. Vesicles
are also produced on the hyphae that grow in the soil. Near the vascular
cylinder, the hyphae branch dichotomously and form large numbers of
thin-walled tubules called arbuscules that invade host cells. The arbus-
cules cause the host membranes to fold inward, creating a plant-fungus in-
terface that has a very large surface area. The arbuscules last for about four-
teen days before they break down on their own or are digested by the host
cell. Host cells whose fungal arbuscules have broken down may be re-
invaded by other hyphae.
Darrell L. Ray
See also: Coevolution; Communities: ecosystem interactions; Commu-

nities: structure; Lichens; Old-growth forests; Symbiosis.

Deacon, J. W. Modern Mycology. 3d ed. Malden, Mass.: Blackwell Science,
1997.
Harley, J. L., and S. E. Smith. Mycorrhizal Symbiosis. New York: Academic
Press, 1983.
Raven, Peter H., Ray F. Evert, and Susan E. Eichhorn. Biology of Plants.
6th ed. New York: W. H. Freeman/Worth, 1999.
427
NATURAL SELECTION
Types of ecology: Evolutionary ecology; Speciation
Natural selection is the process of differential survival and reproduction of indi-

viduals resulting in long-term changes in the characteristics of species. This pro-
cess is central to evolution.
N atural selection is a three-part process. First, there must exist differ-

ences among individuals in some trait. Second, the trait differences
must lead to differences in survival and reproduction. Third, the trait dif-
ferences must have a genetic basis. Natural selection results in long-term
changes in the characteristics of a population. As one of the central pro-
cesses responsible for evolution, natural selection results in both fine-
tuning adaptations of populations and species to their environments and
creating differences among species.
The importance of natural selection was first recognized by Charles
Darwin as the primary mechanism for evolutionary change. Processes that
support natural selection include genetic drift and migration. These pro-
cesses interact with the processes responsible for producing variation (mu-
tation and development) and those responsible for determining the rate
and direction of evolution (mating system, population size, and long-term
ecological changes) to establish the evolutionary path of a species.
The Basic Process

Natural selection occurs through the interaction of three conditions: varia-
tion among individuals in a population in some trait, differences in those
individuals’ fitness as a result of the variations in that trait, and heritable
variation in that trait. If those three conditions are met, then the character-
istics of the population with respect to that trait will change from one gen-
eration to the next until equilibrium with other processes is reached.
An example that demonstrates this process involves the peppered moth.
It has two forms in the United Kingdom, a light-colored form and a dark-
colored form; there is variation in color among individuals. Genetic analysis
has shown that this difference in color is caused by a single gene; the varia-
tion has a heritable basis. The moth is eaten by birds that find their food
by sight. The light-colored form cannot be seen when sitting on lichen-
covered trees, while the dark-colored form can be seen easily. Air pollution
kills the lichen, however, and turns the trees dark in color. Then, the dark-
colored form is hidden and the light-colored form visible. Thus, differ-
428
Natural selection
ences in color lead to fitness differences: Dark moths will become more fit if
air pollution increases, renders them less visible to predators, and thereby
makes it likelier that their genetic material will pass into another genera-
tion. In fact, this proved to be the case: In the early nineteenth century, the
dark-colored form was very rare. In the last half of the nineteenth century,
however, air pollution increased, and the dark-colored form became much
more frequent as a result of natural selection.
Directional Selection
The characteristics of a population can be changed by natural selection in
several ways. If individuals in a population with an extreme value for a
trait have the greatest fitness on average, then the mean value of the trait
will change in a consistent direction, which is called directional selection.
For example, the soil in the vicinity of mines contains heavy metals that are
toxic to plants. Individuals with the greatest resistance to heavy metals
have the highest survivorship. Evolution leads to an increase in resistance.
Stabilizing Selection
If individuals in a population with intermediate values for a trait have the
greatest fitness on average, then the variation in the trait will be reduced,
which is called stabilizing selection. For example, in many species of birds,
individuals with intermediate numbers of offspring have the greatest fit-
ness. If an individual has a small number of offspring, that parent has re-
duced reproduction and a low fitness. If the number of offspring is large,
the parent will not be able to provide enough food for all the young, and
most, or all, will starve, again resulting in reduced reproduction and a low
fitness. Evolution leads to all birds producing the same, intermediate num-
ber of offspring.
Disruptive Selection
If individuals in a population with different values for a trait have the
greatest fitnesses on average and intermediates have low fitness, then the
variation in the trait will be increased. This is called disruptive selection.
For example, for Darwin’s finches, individuals with long, thin bills are able
to probe into rotting cacti to find insects. Individuals with short, thick bills
are able to crack hard seeds. Individuals with intermediate-shaped bills are
not able to do either well and have reduced fitness relative to the more ex-
treme types. Evolution therefore leads to two different species of finch
with different bills.
Natural selection is a slow process. The rate of evolution—that is, re-
sponse to selection—is determined by the magnitude of fitness differences
429
Natural selection
among individuals and the heritability of traits. Fitness differences tend to

be small so that more fit individuals on average may have only a few more
offspring than less fit individuals. Heritabilities of most traits are low to in-
termediate, meaning that most differences among individuals are not a re-
sult of genetic differences. Therefore, even if one individual has many off-
spring and another has few offspring, they may not differ genetically and
no change will occur. For example, if all the beetles in a population were
between one and two centimeters in length and there was selection for
larger beetles, it could take five hundred generations before all beetles
were larger than two centimeters. Also, the direction of selection may
change from one generation to the next, so that no net change occurs.
Correlational Selection
Natural selection does not act on traits in isolation. How a trait affects fit-
ness in combination with other traits—called correlational selection—is
important. For example, fruit flies lay their eggs in rotting fruit. Con-
sidered in isolation, a female should always lay as many eggs as possible.
One fruit is not big enough for all the eggs she might lay, however, so she
must fly from fruit to fruit. Flying requires energy, and the more energy
that is used in flight the less that can be used to make eggs. Hence, natural
selection results in the division of energy between eggs and flight that
yields the greatest overall number of offspring. This example demon-
strates that the result of natural selection is often a trade-off among differ-
ent traits.
Sexual Selection
By acting differently on males and females, natural selection results in sex-
ual selection. This form of selection can explain differences in the forms of
males and females of a species. In general, because male gametes, sperm,
are much smaller and “cheaper” to produce than female gametes, eggs,
more sperm than eggs are produced. As a result, it is possible for one male
to fertilize many eggs, while other males fertilize few or no eggs. For exam-
ple, a lion pride usually consists of one or a few males and many females.
Other males are excluded, and they live separately; larger males are able to
chase away smaller males. The thick mane on male lions helps to protect
their throats when they fight other males. Thus, larger males with thicker
manes tend to survive, fathering more cubs than other males, leading to
additional bias in selection favoring these traits. This is an example of sex-
ual selection because the only selection pressure for the trait in question is
on males; all females, regardless of size, will mate. The result is that males
are larger than females and have manes.
430
Natural selection
Group and Kin Selection

Natural selection can occur not only among individuals but also among
groups. This process is generally known as group selection; when the
groups are composed of related individuals, it is called kin selection.
Group selection operates the same way as individual selection. The same
three conditions are necessary: variation among groups in some trait, fit-
ness differences among groups because of that trait, and a heritable basis
for that trait.
For example, in Australia, rabbits introduced from Europe in 1859
spread rapidly during the next sixty years. In order to control the rabbits, a
virus was introduced in 1950. At first, the virus was very virulent, killing
almost all infected animals within a few days. After ten years, however, the
virus had evolved to become more benign, with infected rabbits living lon-
ger or not becoming sick at all. Virulent strains of the virus grow and repro-
duce faster than benign strains. Therefore, within a single rabbit, virulent
strains have a higher fitness than benign strains. The longer a rabbit lives,
however, the more opportunity there is for the virus to be passed to other
rabbits. Thus, a group of benign viruses infecting a rabbit are more likely to
be passed on than a group of virulent viruses. In this example, group selec-
tion among rabbits resulted in evolution opposite to individual selection
within rabbits; however, group selection and individual selection can re-
sult in evolution in the same direction. In general, natural selection can act
at many levels: the gene, the chromosome, the individual, a group of indi-
viduals, the population, or the species.
Measuring Natural Selection

Natural selection is investigated in two ways: through indirect measure-
ment and through direct measurement. Indirect methods involve observ-
ing the outcome of natural selection and inferring its presence. Direct
methods involve measuring the three parts of the process and following
the course of evolution. Although the direct methods are preferred, as they
provide direct proof of natural selection, in most instances, only indirect
methods can be used.
Indirect methods involve three kinds of observations. First, compari-
sons are made of trait similarities or differences among populations or spe-
cies living in the same or different areas. For example, many species of ani-
mals living in colder climates have larger bodies than those living in
warmer climates. It is inferred, therefore, that colder climates result in nat-
ural selection for larger bodies. Second, long-term studies are done of
traits, in particular changes in a group in the fossil record. For example,
during the evolution of horses, their food, grasses, became tougher and
431
Natural selection
horses’ teeth became thicker. It is inferred, therefore, that tough grass re-
sulted in natural selection for thicker teeth. Third, comparisons are made
of gene frequencies of natural populations, with predictions from mathe-
matical models. Gene frequencies are measured using various techniques,
including scoring differences in appearance, as with light-colored and
dark-colored moths; using electrophoresis to observe differences in pro-
teins; and determining the sequence of base pairs of deoxyribonucleic acid
(DNA). The models make predictions about expected frequencies in the
presence or absence of selection. Indirect methods are best at revealing
long-term responses to evolution and general processes of natural selec-
tion that affect many species. The indirect methods suffer from the prob-
lem that often many processes will result in similar patterns. So, it must be
assumed that other processes were not operating, or other predictions
must be made to separate the processes.
Direct methods involve two kinds of observation. First, there is obser-
vation of changes in a population following some change in the environ-
ment. There are many types of environmental changes, including human-
made changes, natural disasters, seasonal changes, and introductions of
species into new environments. For example, from the changes in the pep-
pered moth following a change in pollution levels, one can measure the ef-
fects of natural selection. The second type of observation is the direct mea-
surement of fitness differences among individuals with trait differences.
For example, individual animals are tagged at an early age and survival
and reproduction are monitored. Then, statistical techniques are used to
find a relationship between fitness and variation among individuals in
some trait. Alternatively, comparisons of traits are made between groups
of individuals, such as breeding and nonbreeding, adults and juveniles, or
live and dead individuals, again using statistical techniques. For example,
lions that breed are larger than lions that do not breed. Direct methods are
best at revealing the relative importance to natural selection of the three
factors (variation, fitness differences, and heritability). The direct methods
suffer from two limitations. It takes a long time for evolution to occur. So,
although one can measure natural selection, it is often not known if it re-
sults in evolution. Also, for many species, it is impossible or impractical to
mark individuals and follow them through their lives.
Many methods can be used to study natural selection and evolution.
Each method provides information about different parts of the process.
Only through the integration of these methods can the entire process of
evolution be revealed.
Knowledge of natural selection is still growing; many questions pro-
posed by Darwin and others are yet to be answered. It is still not known to
432
Natural selection
what extent organisms are well adapted to their environments or whether

the evolution of the parts of the chromosome that are not translated into
proteins are a result of processes that do not involve natural selection. Of
the many theories of how natural selection works, it is still unknown
which ones are the most important in nature and to what extent evolution
is caused by natural selection at the level of the individual, the group, and
the species.
Scientists and researchers combined their knowledge of natural selection
with a revolutionary breakthrough in molecular biology in the 1950’s: the
discovery of the helical structure of deoxyribonucleic acid (DNA) and sub-
sequent discoveries and developments surrounding recombinant DNA
technology. The resultant manipulation of traits in organisms from crops to
mammals led to an age of genetic engineering. Today, strawberries, soy-
beans, dairy products, and a host of other foods, as well as higher organ-
isms including human tissues, can be genetically manipulated to benefi-
cial, as well as often unintended negative, ends. The addition of a new gene
into an organism will result in natural selection on that gene and change
selection on other genes. For example, certain crop plants genetically mod-
ified to resist herbicides have begun to mix with native species considered
weeds, creating a new problem of “superweeds” resistant to herbicides.
An understanding of natural selection is also critical for conservation
biology. During the twentieth century, the rate at which natural areas are
being destroyed and species are becoming extinct has accelerated tremen-
dously. Conservation biology attempts to stop that destruction and pre-
serve species diversity. For extinction of endangered species to be halted, it
must be understood how natural selection will affect these species given
massive environmental changes. By discovering how evolution is occur-
ring under natural conditions, researchers are learning how to design na-
ture preserves to maintain species.
Samuel M. Scheiner, updated by Christina J. Moose

tions and evolutionary explosions; Gene flow; Genetic drift; Genetically
modified foods; Isolating mechanisms; Nonrandom mating, genetic drift,
and mutation; Paleoecology; Population genetics; Punctuated equilibrium
vs. gradualism; Speciation; Species loss.
433
Natural selection

Avers, Charlotte J. Process and Pattern in Evolution. New York: Oxford Uni-
Bell, Graham. Selection: The Mechanism of Evolution. New York: Chapman
and Hall, 1997.
Brandon, Robert N. Adaptation and Environment. Princeton, N.J.: Princeton
Darwin, Charles. On the Origin of Species by Means of Natural Selection. Lon-
don: J. Murray, 1859.
Endler, John A. Natural Selection in the Wild. Princeton, N.J.: Princeton Uni-
Futuyma, Douglas J. Evolutionary Biology. 3d ed. Sunderland, Mass.:
Gould, Stephen J. The Panda’s Thumb. New York: W. W. Norton, 1980.
Pianka, Eric R. Evolutionary Ecology. Boston: Addison-Wesley, 1999.
Provine, William B. Sewall Wright and Evolutionary Biology. Chicago: Uni-
434
NONRANDOM MATING, GENETIC
DRIFT, AND MUTATION
Types of ecology: Evolutionary ecology; Population ecology
Nonrandom mating, genetic drift, and mutation are three mechanisms, besides nat-
ural selection and migration, that can change the genetic structure of a population.
E volution is a process in which the gene frequencies of a population

change over time, and nonrandom mating, genetic drift, and mutation
are all mechanisms of genetic change in populations. These mechanisms
violate the assumptions of the Hardy-Weinberg model of genetic equilib-
rium by increasing or decreasing the frequency of heterozygote genotypes
in the population.
Nonrandom Mating
Nonrandom mating occurs in a population whenever every individual
does not have an equal chance of mating with any other member of the
population. While many organisms do tend to mate randomly, there are
some common patterns of nonrandom mating. Often, individuals tend to
mate with others nearby, or they may choose mates that are most like them-
selves. When individuals choose mates that are phenotypically similar,
positive assortative mating has occurred. If mates look physically differ-
ent, then it is negative assortative mating. Population geneticists use the
term “assortative” because it means “to separate into groups,” usually in a
pattern that is not random. The terms “positive” and “negative” refer to
the probability that mated pairs have the same phenotype more or less of-
ten than expected by chance. Two color varieties of snow geese (Chen
hyperborea), blue and white, are commonly found breeding in Canada, and
they show positive assortative mating patterns based on color. The geese
tend to mate only with birds of the same color; blue mate with blue and
white with white. Since a bird’s color (phenotype) is determined by the
presence of a dominant blue color allele, matings between similar pheno-
types are also matings between similar genotypes. Matings between simi-
lar genotypes cause the frequency of individuals that are homozygous for
the blue or the white allele to be greater, and the frequency of heterozy-
gotes to be less than if mating were random and in Hardy-Weinberg equi-
librium. Negative assortative mating increases the frequency of hetero-
zygote genotypes in the population and decreases homozygote frequency.
435
Nonrandom mating, genetic drift, and mutation
Assortative mating does not change the frequency of the blue or white al-
leles in the goose population; it simply reorganizes the genetic variation
and shifts the frequency of heterozygotes away from Hardy-Weinberg
equilibrium frequencies.
Inbreeding
Inbreeding is the mating of relatives and is similar to positive assortative
mating because like genotypes mate and result in a high frequency of
homozygotes in the population. In assortative mating, only those genes
that influence mate choice become homozygous, but inbreeding increases
the homozygosity of all the genes. High homozygosity means that many of
the recessive alleles that were masked by the dominant allele in heterozy-
gotes will be expressed in the phenotype. Deleterious or harmful alleles
can remain hidden from selection in the heterozygote, but after one gener-
ation of inbreeding, these deleterious alleles are expressed in a homozy-
gous condition and can substantially reduce viability below normal levels.
Low viability resulting from mating of like genotypes is called inbreeding
depression.
Genetic Drift
Genetic drift, like positive assortative mating, reduces the frequency of
heterozygotes in a population, but with genetic drift, the frequency of al-
leles in a population changes. Nonrandom mating does not change allele
frequency. Genetic drift is sometimes called random genetic drift because
the mechanism of genetic change is random and attributable to chance
events in small populations, such that allele frequencies tend to wander or
drift. Statisticians use the term “sampling effect” to describe observed fluc-
tuations from expected values when only a few samples are chosen, and it
is easy to observe by tossing a coin. A fair coin flipped a hundred times
would be expected to produce approximately fifty heads and fifty tails,
plus or minus a few heads or tails. Yet, if the coin is flipped only four times,
it is not too surprising to get four heads or four tails. The probability of get-
ting either all heads or all tails on four consecutive flips is one out of eight,
but the probability of getting all heads or all tails decreases to much less
than one in a billion as the sample size increases from four to a hundred
tosses. Similarly, it is much easier for nonrandom events to occur in small
populations than in large populations. If a population has two alleles with
equal frequency for a particular trait, then the result of random mating can
be simulated by tossing a coin. The frequency of each allele in the next gen-
eration would be determined by flipping the coin twice for each individ-
ual, since sexually reproducing organisms have two alleles for each trait,
436
and counting the number of heads and tails. In a small population, only a
few gametes, each containing one allele for the trait, will fuse to form zy-
gotes. Chance events can cause the frequencies of alleles in a small popula-
tion to drift randomly from generation to generation; often one allele is lost
from the population.
In small populations with fewer than fifty mating pairs, alleles may be
eliminated in fewer than twenty generations by random genetic drift, leav-
ing only one allele for a particular trait in the population. Thus, all individ-
uals would be homozygous for the remaining allele and genetically identi-
cal. Theoretically, in any finite population random genetic drift will occur,
but it is usually negligible if the population size is greater than a hundred.
Sometimes, disasters or disease may drastically reduce the population
size, causing a bottleneck effect. The bottleneck in population size reduces
genetic variability in a population because there are only a few alleles and
results in random genetic drift. Many islands and new populations are
established by a small group of founders that constitute a nonrandom ge-
netic sample because they have only a fraction of the alleles from the origi-
nal large population. Founder effects and bottleneck effects are phenom-
ena that result in a loss of heterozygosity and decreased genetic variability
because of the chance drift in allele frequency away from Hardy-Weinberg
equilibrium values in small populations.
Mutations
Mutations are any changes in the genetic material that can be passed on to
offspring. Some mutations are changes at a single point in the chromo-
some, while at other times, pieces of genetic material are removed, extra
pieces are added, or pieces are exchanged with other chromosomes. All
these changes could result in the formation of new alleles or could change
one allele into a different allele. The random mistakes in the chromosomes
occur at the molecular level, and only later are the changes in information
or alleles translated into phenotypic differences. Thus, mutation is the ulti-
mate source of genetic variability and is random with respect to the needs
of the organism. Most mutations are lethal and are never expressed, but
nonlethal mutations provide the necessary variation for natural selection.
Even though mutations are very important for evolution, they have only a
small effect on allele and genotype frequencies in populations because mu-
tation rates are relatively low. If an allele makes up 50 percent of the gene
pool and mutates to another allele once for every hundred thousand ga-
metes, it would take two thousand generations to reduce the frequency of
the allele by 2 percent. The net effect of mutations is to increase genetic
variability, but at a very slow rate.
437
Studying Genetic Variability

Population geneticists use a wide variety of laboratory, field, and natural
experiments to investigate genetic variability. Natural experiments are sit-
uations that have developed without a scientist intentionally designing an
experiment, but conditions are such that scientists can test a theory. Re-
searchers have used known pedigrees or ancestral histories of zoo animals
and have found that mortality rates of inbred young are often two to three
times higher than for noninbred young. Population geneticists use pedi-
grees to calculate the probability that two alleles are identical by descent;
this research provides an index of the amount of inbreeding in a popula-
tion.
The study of random genetic drift is usually carried out in the labora-
tory. Scientists often use small organisms that reproduce quickly, such as
fruit flies (Drosophila melanogaster), to conserve space and save time. In a
1956 study of eye color conducted by Peter Buri, after only eighteen gener-
ations and sixteen fruit flies per population, more than half of the 107 pop-
ulations started had only one of the two alleles for eye color.
Mutations are so rare that even fruit flies reproduce too slowly for scien-
tists to study the effects of mutations on populations, even though much is
known about the mechanism of mutation by studying Drosophila. Small
bacterial growth chambers can hold many millions of bacterial cells, and,
since they reproduce quickly, even mutations that occur in only one in a
million cells can be detected. In 1955, it was found that mutation rates were
very low in bacteria until caffeine was added to the growth chamber,
whereupon mutation rates increased tenfold. Any chemical or type of radi-
ation that can cause mutations is called a mutagen. Electrophoresis has
also been a useful tool for the study of nonrandom mating, genetic drift,
and mutations, because allele and genotype frequencies can be determined
from samples of the population and unique alleles can be identified.
The Dangers of Inbreeding

Most governments and religions forbid marriages between close relatives
because matings between first cousins result in a 20 percent decrease in
heterozygosity; for those between brothers and sisters, there is an 80 per-
cent decrease in heterozygosity. The decrease in heterozygosity and ge-
netic variation and increase in homozygote frequency often result in in-
breeding depression because deleterious recessive alleles are expressed.
All inbreeding is not undesirable; many of the prizewinning bulls and pigs
at state fairs have some inbreeding in their pedigrees. Most breeds of dogs
were produced by breeding close relatives so that the offspring would
have particular traits.
438
Zookeepers and others that breed and protect rare and endangered spe-
cies must continually be concerned about the negative effects of both in-
breeding and genetic drift. Most zoos are lucky if they have two or three
pairs of breeding adults, and total population sizes are usually very small
compared to those of natural populations. These conditions mean that in-
breeding may reduce the vigor of the population and genetic drift will re-
duce the diversity of alleles in the population, thus reducing the chances of
survival for the captive species. There is hope for rare and endangered spe-
cies if independent inbred lines are crossed, thus reducing the effects of in-
breeding depression, and if breeding adults from other zoos or popula-
tions are traded occasionally, thus increasing the effective population size.
Mutations are the ultimate source of genetic variation and so are very
important in the study of evolution, but the population-level effects of one
mutation are difficult to study because of the low frequency of natural mu-
tations. Certain nonlethal mutations may have little evolutionary impact
but may be important medically because spontaneous mutations result in
hemophilia or dwarfism (achondroplasia) in more than 3 out of 100,000
cases. As exposure to background radiation and chemical levels increases,
mutation rates are likely to increase, as well as the incidence of mutation-
related diseases.
William R. Bromer
See also: Biodiversity; Gene flow; Genetic diversity; Genetic drift; Pollina-
tion; Population genetics; Speciation; Zoos.

Ayala, Francisco J. Population and Evolutionary Genetics: A Primer. Menlo
Park, Calif.: Benjamin/Cummings, 1982.
Crow, J. F. Basic Concepts in Population, Quantitative, and Evolutionary Genet-
ics. New York: W. H. Freeman, 1986.
Fisher, R. A. The Genetical Theory of Natural Selection. New York: Dover,
1958.
Hartl, Daniel L. A Primer of Population Genetics. 3d ed. Sunderland, Mass.:
Mettler, L. E., T. G. Gregg, and H. E. Schaffer. Population Genetics and Evolu-
tion. 2d ed. Englewood Cliffs, N.J.: Prentice-Hall, 1988.
Real, Leslie A., ed. Ecological Genetics. Princeton, N.J.: Princeton University
Press, 1994.
Wilson, Edward O., and W. H. Bossert. A Primer of Population Biology.
439
NUTRIENT CYCLES
Types of ecology: Ecoenergetics; Ecosystem ecology
Within an ecosystem, nutrients move through biogeochemical cycles. Those cycles

involve chemical exchanges of elements among the earth’s atmosphere, water, liv-
ing organisms, soil, and rocks.
A ll biogeochemical cycles—whether the carbon cycle, the hydrologic

cycle, the nitrogen cycle, the phosphorus cycle, or others—have a
common structure, sharing three basic components: inputs, internal cy-
cling, and outputs.
Input of Nutrients
The input of nutrients to an ecosystem depends on the type of biogeo-
chemical cycle. Nutrients with a gaseous cycle, such as carbon and nitro-
gen, enter an ecosystem from the atmosphere. For example, carbon enters
ecosystems almost solely through photosynthesis, which converts carbon
dioxide to organic carbon compounds. Nitrogen enters ecosystems through
a few pathways including lightning, nitrogen-fixing bacteria, and atmo-
spheric deposition. In agricultural ecosystems, nitrogen fertilization pro-
vides a great amount of nitrogen influx, much larger than by any other in-
flux pathways.
In contrast to carbon and nitrogen with input from the atmosphere, the
input of nutrients such as calcium and phosphorus depends on the weath-
ering of rocks and minerals. Soil characteristics and the process of soil for-
mation have a major influence on processes involved in nutrient release to
recycling pools. Supplementary soil nutrients come from airborne parti-
cles and aerosols, as wet or dry depositions. Such atmospheric deposition
can supply more than half of the input of nutrients to some ecosystems.
The major sources of nutrients for aquatic ecosystems are inputs from
the surrounding land. These inputs can take the forms of drainage water,
detritus and sediment, and precipitation. Flowing aquatic systems are
highly dependent on a steady input of detrital material from the watershed
through which they flow.
Internal Cycling
Internal cycling of nutrients occurs when nutrients are transformed in eco-
systems. Plants take up mineral (mostly inorganic) nutrients from soil
through their roots and incorporate them into living tissues. Nutrients
440
Nutrient cycles
in the living tissues occur in various forms of organic compounds and

perform different functions in terms of physiology and morphology. When
these living tissues reach senescence, the nutrients are usually returned
to the soil in the form of dead organic matter. However, nitrogen can
be reabsorbed from senescent leaves and transferred to other living tissues.
Various microbial decomposers transform the organic nutrients into min-
eral forms through a process called mineralization. The mineralized nu-
trients are once again available to the plants for uptake and incorpo-
ration into new tissues. This process is repeated, forming the internal
cycle of nutrients. Within the internal cycles, the majority of nutrients are
stored in organic forms, either in living tissues or dead organic matter,
whereas mineral nutrients represent a small proportion of the total nutri-
ent pools.
Output of Nutrients
The output of nutrients from an ecosystem represents a loss. Output can
occur in various ways, depending on the nature of a specific biogeo-
chemical cycle. Carbon is
released from ecosystems
to the atmosphere in the
The Nitrogen Cycle form of carbon dioxide via
Nitrogen in the process of respiration
atmosphere by plants, animals, and mi-
croorganisms. Nitrogen is
Nitrogen Denitrification
fixation by bacteria lost to the atmosphere in
gaseous forms of nitrogen,
nitrous oxide, and ammo-
Feeding
nia, mostly as by-products
of microbial activities in
Protein Uptake Nitrates in Protein
in plants by roots the soil in animals soil. Nitrogen is also lost
through leaching from the
Nitrification soil and carried out of eco-
systems by groundwater
flow to streams. Leaching
Death and Nitrites Death and also results in export of car-
decomposition decomposition
bon, phosphorus, and other
Nitrifying nutrients out of ecosystems.
bacteria
Output of nutrients from
ecosystems can also oc-
Ammonia cur through surface flow
of water and soil erosion.
441
Nutrient cycles
However, loss of nutrients from one ecosystem may represent input to

other ecosystems. Output of organic matter from terrestrial ecosystems
constitutes the majority of nutrient input into stream ecosystems. Organic
matter can also be transferred between ecosystems by herbivores. For ex-
ample, moose feeding on aquatic plants can transport nutrients to adjacent
terrestrial ecosystems and deposit them in the form of feces.
Considerable quantities of nutrients are lost permanently from ecosys-
tems by harvesting, especially in farming and logging lands, when bio-
mass is directly removed from ecosystems. Fire usually results in the loss
of large amounts of nutrients. Fire kills vegetation and converts portions of
biomass and organic soil matter to ash. Fire causes loss of nutrients
through volatilization and airborne particulate. After fire, many nutrients
become readily available, and nutrients in ash are subject to rapid mineral-
ization. If not taken up by plants during vegetation recovery, nutrients are
likely to be lost from ecosystems through leaching and erosion.
The Hubbard Brook Example

Nutrient cycling has been studied in several intact ecosystems. One of the
most notable experiments was conducted in the Hubbard Brook experi-
mental forest in New Hampshire. The experimental forest was established
initially for forest hydrology research. Begun in the early 1960’s, one of the
longest-running studies of water and nutrient dynamics of forest ecosys-
tems has been on the Hubbard Brook site. Both water and nutrient concen-
trations in precipitation inputs and stream outputs were regularly moni-
tored, allowing estimations of nutrient balances over the watershed
ecosystems.
One of the major findings from the Hubbard Brook study was that un-
disturbed forests exhibit regularity and predictability in their input-output
balances for water and certain chemical elements. Nitrogen, however,
shows a more complex, but still explicable, pattern of stream concentra-
tions. Losses of nitrates from the control watershed are higher in the dor-
mant season, when biological activity is low. Losses are near zero during
the growing season, when biological demand for nitrogen by plants and
microbes are high. Removal of vegetation in the Hubbard Brook forest had
a marked effect on water and nutrient balances. Summer stream flow dur-
ing the devegetation experiment was nearly four times higher than in the
control watershed. The increase in stream flow, combined with increases in
the concentration of nutrients within the stream, resulted in increases in
loss rates of nitrate much higher than those of undisturbed areas. Similarly,
loss of potassium used in large quantities by plants showed the greatest in-
crease.
442
Nutrient cycles
Nutrient Uptake and Competition

Ecosystem nutrient cycling is critical for plant growth and ecosystem pro-
ductivity. Plant uptake of essential nutrient elements is related to nutrient
availability, root absorption surface, rooting depth, and uptake kinetics of
roots. A nutrient-rich site usually supports more plants of different species
than a site with fewer available nutrients. Nutrient competition among
plants is usually manifested through physiological, morphological, and
ecological traits. Usually grasses and forbs can coexist in one grassland
ecosystem, for example, through different rooting depth. To compete for
less soluble nutrients such as phosphorus, plants usually extend their root
surfaces using symbiotic relationships with mycorrhizae. Differential sea-
sonality in nutrient uptake and rooting depth become more critical to com-
pete for limited nutrients.
Yiqi Luo
See also: Balance of nature; Biomass related to energy; Competition; Food

chains and webs; Geochemical cycles; Herbivores; Hydrologic cycle; Om-
nivores; Phytoplankton; Rain forests and the atmosphere; Trophic levels

Aber, J. D., and J. M. Melillo. Terrestrial Ecosystems. 2d ed. San Diego: Aca-
demic Press, 2001.
Likens, G. E., et al. Biogeochemistry of a Forested Ecosystem. New York:
Springer-Verlag, 1977.
Schlesinger, W. H. Biogeochemistry: An Analysis of Global Change. 2d ed. San
Diego: Academic Press, 1997.
443
OCEAN POLLUTION
AND OIL SPILLS
Types of ecology: Aquatic and marine ecology; Ecotoxicology
Oil spills resulting from human error often affect marine and coastal areas. Past oil
spills in different areas of the world demonstrate that environmental damage de-
pends on the toxicity and the persistence of the oil; both vary widely depending on
a variety of factors.
Why Oil Spills Occur

Almost all imported and Alaskan oil is transported to U.S. refineries and
consumers by oceangoing tankers. Most oil spills result from marine trans-
portation accidents, with human error usually playing a major role. Navi-
gation errors, equipment malfunctions, bad judgment, and even the inabil-
ity of all crew members to speak a common language have all been major
contributing factors in the largest and most environmentally damaging oil
spills. Not all oil spills are environmental disasters, but spilled oil can deci-
mate plant and animal populations by a combination of mechanical toxic-
ity and chemical toxic effects resulting from an organism’s physiological
reaction to the chemicals present in oil.
Environmental Damage
The most common sight during an oil spill is dark, gelatinous masses of
“mousse”—an oil and water emulsion that floats on the water, sticking to
everything with which it comes into contact. Mousse usually causes the
majority of the environmental damage during an oil spill by the process of
mechanical toxicity, as it suffocates and smothers organisms that ingest it
or are covered by it. Seabirds and furry marine mammals are highly sus-
ceptible to this process, succumbing to exposure, dehydration, or starva-
tion.
Crude oil is a complex mixture of thousands of different chemicals
called hydrocarbons, named after a molecular structure based on hydro-
gen and carbon atoms. Different hydrocarbons vary in their chemical
properties, toxicity, and behavior during an oil spill. The major groups are
classified by molecular geometry and weight. The low-molecular-weight
molecules (aliphatics) are single-bonded, chain-shaped molecules, such as
gasoline. They are the most chemically reactive and volatile, and they are
acutely toxic. These compounds tend to evaporate or burn easily during
444
Ocean pollution and oil spills
an oil spill and therefore do not persist in the environment for long peri-
ods. Intermediate-molecular-weight hydrocarbons, or aromatics, are ring-
shaped molecules, such as benzene. They are also highly reactive and
cause biological impacts because of both acute and chronic toxicity. Aro-
matic hydrocarbon compounds are more environmentally persistent than
aliphatics. Since many are carcinogens, they can cause different forms of
biological damage, disease, and death even after a long time period and
in low doses. The high-molecular-weight oil compounds are mostly poly-
cyclic aromatic hydrocarbons structured of ring shapes bonded together to
form molecules. Although they are not very chemically reactive and do not
dissolve well in water, many are carcinogenic. They tend to be very envi-
ronmentally persistent.
For hundreds of millions of years before humans evolved, oil was
“spilled” naturally into the world’s oceans by natural oil seeps—fractures
in the earth’s crust that tap deep, oil-bearing rocks. A variety of natural
processes act to reduce the environmental impacts of this oil, and these
same processes also take place during a human-caused oil spill. Oil is dis-
persed from the oil slick and into the larger environment by five basic pro-
cesses. Evaporation of the low-molecular-weight hydrocarbon com-
pounds removes most of the oil relatively quickly. Sunlight can degrade
additional oil in a process called photodegradation if the oil is exposed for
enough time. Because oil is an organic substance, additional oil is removed
by natural biodegradation thanks to “oil-eating” microorganisms. Most of
the rest of the oil either washes up onto a coastal area or breaks up into
heavy “tar balls” rich in high-molecular-weight hydrocarbons that eventu-
ally sink.
Some oil spills put so much oil into the environment that these pro-
cesses cannot respond quickly enough to prevent environmental damage.
Other factors can also enhance environmental damage from oil spills.
Some types of oil or refined petroleum products are more toxic than others.
Oil spills in cold climates generally cause more damage because cold tem-
peratures retard evaporation and the microbial metabolic rates necessary
for rapid oil removal. Furthermore, sunlight is often of low intensity, which
retards photodegradation. Wave conditions and tidal currents can affect
how much oil washes up onto a coastal area and how rapidly it is moved
elsewhere or removed. Finally, the amount of environmental damage from
an oil spill is highly dependent on the type of coastal environment oiled in
the spill, as coastal environments vary in density (or biomass) and varieties
of wildlife. Coastlines also vary in the degree to which they are sheltered
from natural oil-removal processes. In general, rocky headlands, wave-cut
rock platforms, and reefs exposed to high wave activity suffer far less dam-
445
age during an oil spill than do sheltered marshes, tidal flats, and mangrove
forests. The damage on beaches is related to the grain size of the beach sed-
iment. Fine-sand beaches are relatively flat and hard-packed, and oil does
not soak into the sediment or persist for long. Oil will soak deeply into
coarse sand, gravel, and shell beaches, causing more damage over a longer
period.
Most of what has been learned about oil spill behavior, environmental
damage, and oil spill cleanup techniques comes from studying past spills.
In most cases, spill prevention is far cheaper and more effective than spill
response, and cleanup efforts usually capture very little of the spilled oil.
Ixtoc I
The Ixtoc I spill of June 3, 1979, was the result of an explosion, or “blow-
out,” of an offshore oil well that was drilling into a subsurface oil reservoir.
Although human error was definitely a factor, the cause of the blowout re-
mains unresolved. It has been blamed on the use of drilling mud that was
not dense enough to counteract the pressure of the oil and gas at depth, as
well as on the improper installation of the blowout preventer—a fail-safe
device used on drilling rigs to prevent just this type of disaster. The result
was a continuous 290-day oil spill, during which an estimated 475,000 met-
ric tons of crude oil (one metric ton equals approximately five barrels) were
released into the environment. In addition to doing considerable environ-
mental damage on the coast of Mexico, oil fouled much of the barrier is-
land coast of Texas. However, most of the oil did not make it to shore, and
the final accounting for this spill gives a good indication of the long-term
fate of spilled oil in offshore areas: 1 percent burned at the spill site, 50 per-
cent evaporated, 13 percent photodegraded or biodegraded, 7 percent
washed up on the coast (6 percent in Mexico, 1 percent in Texas), 5 percent
was mechanically removed by skimmers and booms, and 24 percent sank
to the seafloor (assumed by mass balance).
The Exxon Valdez

The Exxon Valdez oil spill—which occurred in Prince William Sound, Alaska,
on March 24, 1989—is a good example of how environmental damage fol-
lows human error and inadequate response. After departing Port Valdez
with a full cargo, the Exxon Valdez oil tanker struck a well-charted sub-
merged rock reef located 1.6 kilometers outside the shipping lane. The ship
was under the command of an unlicensed third mate in calm seas and left
the shipping lane with permission from the Coast Guard to avoid ice. How-
ever, it strayed too close to the reef before evasive action was attempted.
The captain was under the influence of alcohol during events leading to the
446
Seabirds are often casualties of ocean oil spills, succumbing to exposure, dehydra-
tion, starvation, suffocation, or the oil’s toxicity. In 1989, experts estimated that,
in addition to thousands of sea otters and deer, more than half a million seabirds
died as a result of the Exxon Valdez oil spill in Prince William Sound, Alaska.
(PhotoDisc)
accident. His blood alcohol level nine hours after the grounding was mea-
sured at 0.06 percent; the estimate at the time of the accident was 0.19 per-
cent—almost twice the legal level for drivers in California. Convicted of
negligence and stripped of his commander’s license, he was subsequently
employed as an instructor to teach others to operate supertankers.
Leaking oil was observed immediately. Oil-spill response crews funded
by Exxon and the Alyeska Pipeline Consortium—oil companies that used
the Port Valdez terminal—were poorly prepared and reacted too slowly
and with inadequate equipment. The first response arrived ten hours after
the accident with insufficient booms and skimmers. Chemical dispersants
applied to break up the oil slick were ineffective in the calm seas and
caused the oil slick to thin and spread more rapidly. Four days later, the
weather changed: 114-kilometer-per-hour winds mixed the oil with seawa-
ter, creating a frothy mousse. More than 65,000 metric tons of oil spilled
over several weeks. About 15,600 square kilometers of ocean and 1,300 ki-
lometers of shoreline were affected.
Federal estimates of wildlife mortality include 3,500 to 5,500 otters;
580,000 seabirds; and 300 deer poisoned by eating oiled kelp. Economic
damages totaled more than $5 billion. The long-term effects on commercial
447
marine organisms, larval organisms, and bottom-dwelling life are not

known.
Exxon promised to clean nearly 500 kilometers of shoreline by Septem-
ber, 1989, but cleaned only 2 kilometers during the first month after the
spill. Exxon and its contractors used a variety of cleanup techniques, in-
cluding placing booms and skimmers, sopping up oil with absorbent
materials, scraping oil by hand from rocks, stimulating the growth of oil-
eating bacteria cultures, and washing coastal areas with cold water, hot
water, and steam. The use of hot water and steam was effective at cosmetic-
ally removing surface oil, but it did not remove oil that had soaked into the
sediment; the technique killed most organisms that had escaped the oil.
The oil washed from the beach was to be collected by booms and skimmers
offshore, but this process was so inefficient that much of the oil migrated to
tide pools that had not been affected by the spill directly. Ironically, only
eighteen months after the spill, life had returned to oiled coasts that had re-
ceived little or no cleanup, while beaches cleaned with hot water were still
relatively sterile and required several years to repopulate. Exxon an-
nounced that it would not return to clean more shoreline in 1990 but re-
lented under threat of a court order from the Coast Guard to enforce fed-
eral cleanup requirements. During the summer of 1990, shoreline cleanup
resumed, including application of fertilizer to stimulate growth of natu-
rally occurring oil-eating bacteria, a technique that is not very efficient in
the cold waters of southern Alaska.
The tale of the Exxon Valdez is not complete without mentioning that the
Port Valdez Coast Guard did not have state-of-the-art radar equipment for
monitoring ship movement in this heavily used and environmentally sen-
sitive area. In the early 1980’s, federal and state funds for monitoring the
Port Valdez oil companies’ compliance with oil-spill preparedness legisla-
tion had been cut by more than 50 percent. The original environmental im-
pact statement for oil-handling activity in Prince William Sound included
an agreement that defines cleanup responsibility for oil spills. Exxon, as
the company responsible for the spill, was to pay the first $14 million of
cleanup costs, with $86 million in additional cleanup funds from the
Alyeska contingency fund. Thus, the maximum financial responsibility to
oil companies from a spill was $100 million unless the spill was judged to
be caused by negligence. Cleanup activities ceased eighteen months after
the spill with total expenditure of $2.2 billion; most of this at taxpayer ex-
pense. In 1994, a federal court unanimously awarded $5.3 billion in puni-
tive and compensatory damages, the largest-ever jury award, to some
thirty-five thousand people impacted by the spill. By June, 1999, Exxon
had yet to pay a single dollar as the case continued through the legal pro-
448
cess. Finally, it is interesting to note that Exxon’s estimate of cleanup costs

in late 1989 were $500 million, and it carried $400 million of oil spill liabil-
ity insurance. Exxon saved $22 million by not building the Exxon Valdez
with a double hull; its 1988 annual profits were $5,300 million.
According to National Oceanic and Atmospheric Administration (NOAA)
estimates, less than 1 percent of Exxon Valdez’s oil burned at the site,
20 percent evaporated, 8 percent was mechanically removed, and nearly
72 percent was deposited on the seafloor. According to Exxon’s esti-
mates, 7 percent of the oil burned at the site, 32 percent evaporated, 9 per-
cent photodegraded or biodegraded, 15 percent was mechanically re-
moved, and 37 percent was assumed deposited on the seafloor.
Operation Desert Storm

The February, 1991, Operation Desert Storm oil spill—the largest oil spill in
history—occurred when the Iraqi military opened valves and pumps at Sea
Island Terminal, a tanker loading dock located 16 kilometers off the coast of
Kuwait. This facility has a production capacity of 100,000 barrels per day,
about three Exxon Valdez loads each week. The Iraqis also opened plugs on
five Kuwaiti tankers, spilling an additional 60,000 barrels. The estimate for
the entire spill is 6 million barrels, or roughly 30 times the volume of the Ex-
xon Valdez. About 650 square kilometers of coast were heavily contaminated.
Three days after starting the spill, the Iraqis ignited the oil leaking from
the terminal. This was the best thing to happen from an environmental per-
spective. During most spills, more oil is removed by natural evaporation
than by any cleanup technique; igniting the oil merely speeds up this pro-
cess. Burning can be an important mechanism for removing oil from the
sea and avoiding environmental damage, and tests have shown purpose-
ful ignition in open water away from the coast to be an excellent oil-slick
fighting strategy. However, this must be done within the first few hours of
the spill—in order to maintain the fire, the slick must be more than 1 milli-
meter thick and must contain relatively little emulsified water. To maintain
thickness, the slick is best surrounded with fireproof booms. However, at
the time of the Operation Desert Storm spill, almost all the fireproof booms
in the world were in Prince William Sound. Saudi Arabia also used dis-
persants on portions of the slick, but this effort was too late to be effective
before a thick mousse had formed.
The prime objectives of causing the spill were to hamper an amphibious
military landing by oiling the beaches and to disrupt desalinization of
drinking water at Khafji and Jubail, the two primary sources of potable
water for Saudi Arabia. The Saudis used booms to protect the plant intakes
with great success. The retreating Iraqis also ignited more than seven hun-
449
dred of about one thousand inland wells, resulting in an additional 6 mil-

lion barrels per day burned. This volume eventually made the marine spill
insignificant, and the burning created 3 percent of total global carbon emis-
sions during the time period of the event.
The Arabian Gulf is an unusual body of water. It is very shallow (aver-
age 33 meters) and is nearly enclosed as a marine basin. Because it is also
microtidal (the tidal range is less than 0.6 meter), it flushes out slowly (once
every two hundred years, compared with once every few days for Prince
William Sound). It is also important to remember that this is not a pristine
marine environment. Natural oil seeps are very common, there is a general
lack of environmental standards and poor cooperation among Persian
Gulf nations, and virtually no oil spill preparations or equipment were
present in this part of the world. Earlier spills had occurred in the region,
but they typically were associated with ongoing wars; the hostile environ-
ment made it difficult to utilize spill abatement specialists and equipment.
For example, during the Iran-Iraq War, Iraq attacked an Iranian offshore
platform (Nowruz) in 1982, spilling more than 2 million barrels of oil, a
volume nearly half as large as the Operation Desert Storm spill. Losses of
marine mammals and birds were great, and populations had not re-
bounded by the time of Operation Desert Storm.
During the Operation Desert Storm oil spill, about 180 kilometers of
Saudi Arabian coastline were oiled (65 kilometers were severely dam-
aged), and oil reached south as far as the United Arab Emirates and Bah-
rain. Much of the southern Kuwaiti coast (sea-grass beds, marsh, and man-
groves) was severely damaged, and about 25 percent of the Saudi shrimp
industry was lost. Although some twenty thousand wading birds were
killed, no deaths of dolphins or dugongs were reported. However, these
animals suffered greatly during the Iran-Iraq War. Estimates of the time re-
quired for ecological renewal of the Persian Gulf following the Operation
Desert Storm spill (one to four years) were relatively short for two reasons:
The high water temperature results in high microbial activity and biodeg-
radation of the oil, and much of the oil was burned.
James L. Sadd
See also: Acid deposition; Lakes and limnology; Marine biomes; Pollution
effects; Reefs.

Alaska Wilderness League. Preventing the Next Valdez: Ten Years After Ex-
xon’s Spill New Disasters Threaten Alaska’s Environment. Washington,
D.C.: Alaska Wilderness League, 1999.
450
Etkin, Dagmar Schmidt. Financial Costs of Oil Spills in the United States.
Arlington, Mass.: Cutter Information, 1998.
_______. Marine Spills Worldwide. Arlington, Mass.: Cutter Information,
1999.
Hall, M. J. Crisis on the Coast. Portland, Oreg.: USCG Marine Safety Office,
1999.
Smith, Roland. The Sea Otter Rescue: The Aftermath of an Oil Spill. New York:
Puffin Books, 1999.
451
OLD-GROWTH FORESTS
Types of ecology: Biomes; Ecosystem ecology; Restoration and
conservation ecology
Ancient ecosystems, old-growth forests consist of trees that have never been har-
vested. These forests are, in some cases, the only habitat for a number of plant and
animal species.
T he timber industry views large, old trees as a renewable source of fine

lumber, but environmentalists see them as part of an ancient and
unique ecosystem that can never be replaced. In the 1970’s scientists began
studying the uncut forests of the Pacific Northwest and the plants and ani-
mals that inhabited them. In a U.S. Forest Service publication, Ecological
Characteristics of Old-Growth Douglas-Fir Forests (1981), Forest Service biolo-
gist Jerry Franklin and his colleagues showed that these forests were not
just tangles of dead and dying trees but rather a unique, thriving ecosys-
tem made up of living and dead trees, mammals, insects, and fungi.
Old-Growth Forest Ecosystem

The forest usually referred to as old growth occurs primarily on the west-
ern slope of the Cascade Mountains in southeast Alaska, southern British
Columbia, Washington, Oregon, and Northern California. The weather
there is wet and mild, ideal for the growth of trees such as Douglas fir, ce-
dar, spruce, and hemlock. Studies have shown that there is more biomass,
including living matter and dead trees, per acre in these forests than any-
where else on earth. Trees can be as tall as 300 feet (90 meters) with diame-
ters of 10 feet (3 meters) or more and can live as long as one thousand years.
The forest community grows and changes over time, not reaching biologi-
cal climax until the forest primarily consists of hemlock trees, which are
able to sprout in the shade of the sun-loving Douglas fir.
One of the most important components of the old-growth forest is the
large number of standing dead trees, or snags, and fallen trees, or logs, on
the forest floor and in the streams. The fallen trees rot very slowly, often
taking more than two hundred years to completely decompose. During
this time they are important for water storage, as wildlife habitat, and as
“nurse logs” where new growth can begin. In fact, seedlings of some trees,
such as western hemlock and Sitka spruce, have difficulty competing with
the mosses on the forest floor and need to sprout on the fallen logs.
Another strand in the complex web of the forest consists of mycorrhizal
452
Old-growth forests
fungi (mycorrhizae), which attach themselves to the roots of the trees and
enhance their uptake of water and nutrients. The fruiting bodies of these
fungi are eaten by small mammals such as voles, mice, and chipmunks,
which then spread the spores of the fungi in their droppings. There are nu-
merous species of plants and animal wildlife that appear to be dependent
on this ecosystem to survive.
Protecting the Forest

By the 1970’s most of the trees on timber industry-owned lands had been
cut. Their replanted forests, known as second growth, would not be ready
for harvest for several decades, so the industry became increasingly de-
pendent on public lands for their raw materials. Logging of old growth in
the national forests of western Oregon and Washington increased from 900
million board feet in 1946 to more than 5 billion board feet in 1986.
Environmentalists claimed that only 10 percent of the region’s original
forest remained. Determined to save what was left, they encouraged the
use of the evocative term “ancient forest” to counteract the somewhat neg-
ative connotations of “old growth.” Then they were given an effective tool
in the northern spotted owl. This small bird was found to be dependent on
Old-growth forests in the Pacific Northwest are the only habitat for not only the fa-
mous endangered northern spotted owl but also a host of other wildlife species.
(PhotoDisc)
453
Old-growth forests
old growth, and its listing under the federal Endangered Species Act in
1990 caused a decade of scientific, political, and legal conflict.
Under law, the U.S. Forest Service was required to protect enough of the
owl’s habitat to ensure its survival. An early government report identified
7.7 million acres of forest to be protected for the bird. Later, the U.S. Fish
and Wildlife Service recommended 11 million acres. In 1991 U.S. District
Court judge William Dwyer placed an injunction on all logging in spotted
owl habitat until a comprehensive plan could be finalized. The timber in-
dustry responded with a prediction of tens of thousands of lost jobs and re-
gional economic disaster. In 1993 President Bill Clinton convened the For-
est Summit conference in Portland, Oregon, to work out a solution. The
Clinton administration’s plan, though approved by Judge Dwyer, satisfied
neither the industry nor the environmentalists, and protests, lawsuits, and
legislative battles continued.
As the twentieth century came to an end, timber harvest levels had been
significantly reduced, the Northwest’s economy had survived, and addi-
tional values for old-growth forests were found: habitat for endangered
salmon and other fish, a source for medicinal plants, and a repository for
benefits yet to be discovered. The decades-long controversy over the for-
ests of the Northwest had a deep impact on environmental science as well
as natural resource policy and encouraged new interest in other native for-
ests around the world, from Brazil to Malaysia to Russia.
Joseph W. Hinton
See also: Endangered animal species; Forests; Habitats and biomes; Lakes
and limnology; Mountain ecosystems; Rain forests; Rain forests and the at-
mosphere; Savannas and deciduous tropical forests; Slash-and-burn agri-
culture; Taiga; Tundra and high-altitude biomes.

Dietrich, William. The Final Forest: The Battle for the Last Great Trees of the Pa-
cific Northwest. New York: Penguin Books, 1993.
Durbin, Kathie. Tree Huggers: Victory, Defeat, and Renewal in the Northwest
Ancient Forest Campaign. Seattle: Mountaineers, 1996.
Kelly, David, and Gary Braasch. Secrets of the Old Growth Forest. Salt Lake
City, Utah: Gibbs-Smith, 1988.
Maser, Chris. Forest Primeval: The Natural History of an Ancient Forest.
Corvallis: Oregon State University Press, 2001.
454
OMNIVORES
Types of ecology: Behavioral ecology; Ecoenergetics
Omnivores are animals that eat both plants and animals. They are found in all
types of animals, including arthropods, fish, birds, and mammals. Omnivore diets
may vary seasonally.
M any animals are either herbivores, which eat only plant food, or car-
nivores, which eat only the flesh of other animals. The preference
for one type of food or the other depends largely on the type of digestive
system that the animal has, and the resources it can put into its “energy
budget.”
Ecological Advantages
Meat is generally easier to digest and requires a less complex digestive sys-
tem and a relatively short intestinal tract. However, in order to get meat,
carnivores have to invest a lot of time hunting their prey, and the outcome
of a hunt is always uncertain. The food of herbivores is much easier to ob-
tain, since plants do not move and all the herbivore has to do is graze on
the grasses, leaves, or algae readily available around it. However, the cellu-
lose that plants are made of is very tough to digest, and thus herbivores
must have a much lengthier and more complex digestive tract than carni-
vores. Many herbivores are ruminants, with multipart stomachs, which
have to chew and digest their food more than once in order to get adequate
nutrition from it.
Carnivores and herbivores are also vulnerable to a loss of their food
source. Herbivores whose digestive systems are specialized to process
only one type of food will starve if that food becomes scarce as a result of
drought or some other climatic change. Carnivores often have specialized
hunting patterns that cannot be changed if the prey (usually herbivores)
become scarce due to loss of their own food source.
Omnivores maximize their ability to obtain food by having digestive
tracts capable of processing both plant and animal food, although they are
usually not capable of digesting the very tough plant material, such as
grasses and leaves, that many large herbivores eat. Omnivores may also be
scavengers, eating whatever carrion they may come across. Omnivores of-
ten lack the specialized food-gathering ability characteristic of pure carni-
vores and herbivores. Many animals often thought of as carnivores are ac-
tually omnivores, eating both plants and animals.
455
Omnivores
Diversity of Omnivores
Omnivores can be found among all types of animals, living on land and in
water. They include fish, mollusks, arthropods, birds, and mammals.
Most insects are either herbivores, such as grasshoppers, or carnivores
such as mantises. However some, such as yellow jacket wasps, are omni-
vores, eating other insects, fruit, and nectar. Omnivorous snails and slugs
eat algae, leaves, lichens, insects, and decaying plant and animal matter.
Their main organ for eating is called a radula, a tonguelike, toothed organ
that is drawn along rocks, leaves, or plants to scrape off food; it is also used
to bore holes through shells of other mollusks, to get to their flesh.
Omnivorous fish include the common carp, goldfish, catfish, eels, and
minnows. Since a fish’s food is often suspended in the medium through
which the fish swims—water—being able to gulp up whatever comes into
its mouth is an efficient way for a fish to eat. Similarly, bottom-feeders (fish
that suck up material from the floor of whatever body of water they in-
habit) also benefit from not needing to sort through the material before
they ingest it.
Many birds are omnivores, such as robins, ostriches, and flamingos. The
pink or red color of flamingos occurs because they eat blue-green algae and
higher plants which contain the same substances that make tomatoes red.
They also eat shrimp and small mollusks.
Mammal omnivores include bears; members of the weasel family, such
as skunks; the raccoon family (raccoons and coatimundis); monkeys; apes;
and humans. Raccoons and coatis, found only in the Americas, eat insects,
crayfish, crabs, fish, amphibians, birds, small mammals, nuts, fruits, roots,
and plants. Like other omnivores, they also eat carrion. Bears eat grass,
roots, fruits, insects, fish, small or large mammals, and carrion.
Sanford S. Singer
See also: Balance of nature; Biomass related to energy; Food chains and
webs; Herbivores; Nutrient cycles; Predation; Trophic levels and ecological
niches.

Kay, Ian. Introduction to Animal Physiology. New York: Springer-Verlag,
1999.
Lauber, Patricia. Who Eats What? New York: HarperTrophy, 1995.
Llamas, Andreu. Crustaceans: Armored Omnivores. Milwaukee: Gareth Ste-
vens, 1996.
McGinty, Alice B. Omnivores in the Food Chain. Logan, Iowa: Powerkids
Press, 2002.
456
OZONE DEPLETION AND
OZONE HOLES
Types of ecology: Ecotoxicology; Global ecology
Ozone occurs naturally in the atmosphere and absorbs ultraviolet radiation from
the sun. In the past few decades, a “hole” in the atmosphere’s ozone layer has been
recorded over Antarctica, and its size, although fluctuating, has increased over
time. Scientists are concerned with the damage to all living organisms from ultra-
violet radiation if the atmosphere’s ozone continues to decrease.
Ozone in the Atmosphere

Ozone, although only a minor component of the atmosphere, plays a vital
role in the survival of life on earth. Ozone molecules absorb high-energy
ultraviolet (UV) radiation, which humans perceive as light, from the sun.
Absorption of ultraviolet radiation in the ozone layer, a region of the
stratosphere that contains the maximum concentration of ozone, prevents
most such light from reaching the surface of the planet. If none of the sun’s
ultraviolet radiation were blocked by the ozone layer, it would be difficult
for most forms of life, including humans, to survive on land.
The concentration of ozone in the atmosphere is highly variable, chang-
ing with altitude, geographic location, time of day, time of year, and pre-
vailing local atmospheric conditions. Long-term fluctuations in ozone con-
centration are also seen, some of which are related to the solar sunspot
cycle. While long-term average ozone concentrations are relatively stable,
short-term fluctuations of as much as 10 percent in total column abun-
dance of ozone as a result of the natural variability in ozone concentration
are often observed.
Discovery of a “Hole”
Beginning in the early 1970’s, a new and unexpected decrease in strato-
spheric ozone concentration was first observed. The decrease was local-
ized in geography to the Southern Polar region, and in time to early spring
(which begins in October in the Southern Hemisphere). The initial de-
crease in ozone was small, but by 1980, decreases in total column abun-
dance of ozone of as much as 30 percent were being seen, well outside the
range of variation expected as a result of random fluctuations. This sea-
sonal destruction of stratospheric ozone above Antarctica, which by 1990
had reached 50 percent of the total column abundance of ozone, was soon
given the label “ozone hole.”
457
Ozone depletion and ozone holes
Average Size of the Ozone Hole, 1980-2000

30
25 Area of North America

Size (million square kilometers)
20
15
Area of Antarctica
10
5 Ozone values < 220 Dobson units

Average area - 30 day maximum
Vertical lines = minimum & maximum area
0
1980 1985 1990 1995 2000
Year
Source: Goddard Space Flight Center. National Aeronautics and Space Administration.
http:\\toms.gsfc.nasa.gov\multi\oz_hole_area.jpg; accessed April 15, 2002.
The Role of CFCs

While it was initially unclear whether formation of the Antarctic ozone
hole stemmed from natural causes or from anthropogenic effects on the en-
vironment, extensive field studies combined with the results of laboratory
experiments and computer modeling of the atmosphere quickly led to a
consistent and detailed explanation for ozone-hole formation. The forma-
tion of the ozone hole has two principal causes: chemical reactions that oc-
cur generally throughout the stratosphere, and special conditions that ex-
ist in the Antarctic region.
Under normal conditions, the concentration of ozone in the strato-
sphere is determined by a balance between reactions that remove ozone
and those that produce ozone. The removal reactions are mainly cata-
lytic chain reactions, in which trace atmospheric chemical species destroy
ozone molecules without themselves being consumed. In such processes,
it is possible for one chain carrier to remove many ozone molecules before
being itself removed. The trace species involved in ozone removal include
hydrogen oxides and nitrogen oxides, formed primarily by naturally oc-
458
curring processes, and chlorine and bromine atoms and their correspond-
ing oxides.
A major source of chlorine in the stratosphere is the decomposition of
a class of compounds called chlorofluorocarbons (CFCs). Such compounds
can be used in refrigeration and air conditioning, as aerosol propellants,
and as solvents. Chlorofluorocarbons are extremely stable in the lower at-
mosphere, with lifetimes of several decades. The main fate of chlorofluoro-
carbons is slow migration into the stratosphere, where they absorb ultravi-
olet light and release chlorine atoms. The chlorine atoms produced from
the breakdown of chlorofluorocarbons in the stratosphere provide an ad-
ditional catalytic process by which stratospheric ozone can be destroyed.
A similar set of reactions involving a class of bromine-containing com-
pounds called halons, used in some types of fire extinguishers, leads to ad-
ditional ozone destruction. By 1986, the average global loss of strato-
spheric ozone caused by the release of chlorofluorocarbons, halons, and
related compounds into the environment was estimated to be 2 percent.
The Antarctic Stratosphere

While the decomposition and subsequent reaction of chlorofluorocarbons
and other synthetic compounds explains the small general decline in
ozone concentration observed in the stratosphere, additional processes are
needed to account for the more massive seasonal ozone depletion ob-
served above Antarctica. These processes involve a set of special condi-
tions that in combination are unique to the stratosphere above Antarctica.
During daylight hours, a portion of the chlorine present in the strato-
sphere is tied up in the form of reservoir species, compounds such as hy-
drogen chloride and chlorine nitrate that do not react with ozone. This
slows the rate of removal of ozone by chlorine. Processes that directly or in-
directly involve absorption of sunlight transform reservoir species into
ozone-destroying chlorine atoms. During the Antarctic winter, when sun-
light is entirely absent, stratospheric chlorine is rapidly converted into res-
ervoir species.
In the absence of additional chemical processes, the onset of spring in
Antarctica and the return of sunlight would convert a portion of the reser-
voir compounds into reactive chlorine species and reestablish the balance
between ozone-producing and ozone-destroying processes. However, the
extremely low temperatures occurring in the stratosphere above Antarc-
tica during the winter months leads to the formation of polar stratospheric
clouds, which, because of the extremely low concentration of water vapor
in the stratosphere, do not form during other seasons or outside the polar
regions of the globe. The ice crystals that compose the clouds act as cata-
459
lysts that convert reservoir species into diatomic chlorine and other gas-
eous chlorine compounds that, in the presence of sunlight, re-form ozone-
destroying species. At the same time, the nitrogen oxides found in the res-
ervoir species are converted into nitric acid, which remains attached to the
ice crystals. As these ice crystals are slowly removed from the stratosphere
by gravity, the potential for conversion of active forms of chlorine into res-
ervoir species is greatly reduced. Because of this, when spring arrives,
large amounts of ozone-destroying chlorine species are produced by the
action of sunlight, and only a small fraction of this reactive chlorine is con-
verted into reservoir species. The increased rate of ozone removal caused
by the abundance of reactive chlorine present in the stratosphere leads to
ozone depletion and formation of the ozone hole.
An additional process important in formation of the ozone hole is the
unique air-circulation pattern in the stratosphere above Antarctica. During
the winter and early spring, a vortex of winds circulates about the South
Pole. This polar vortex minimizes movement of ozone and reservoir-form-
ing compounds from other regions of the stratosphere. As this polar vortex
breaks up in midspring, ozone concentrations in the Antarctic stratosphere
return to normal levels, and the ozone hole gradually disappears.
Study and Interpretation

Researchers utilize a great diversity of devices and techniques in their
study and interpretation of atmospheric ozone. One popular technique is
the use of models. A good model is one that simulates the interrelation-
ships and interactions of the various parts of the known system. The weak-
ness of models is that, often, not enough is known to give an accurate pic-
ture of the total system or to make accurate predictions. Most modeling is
done on computers. Scientists estimate how fast chemicals such as CFCs
and nitrous oxide will be produced in the future and build a computer
model of the way these chemicals react with ozone and with one another.
From this model, it is possible to estimate future ozone levels at different
altitudes and at different future dates.
Arctic Depletion?
Similar processes appear to be at work in the Arctic stratosphere, leading
to ozone depletion, as in the Antarctic; however, the National Oceanic and
Atmospheric Administration (NOAA) Aeronomy Laboratory in Boulder,
Colorado, reported a discrepancy between observed ozone depletion and
predicted levels, based on models that account accurately for the Antarctic
depletions. This report suggests that some other mechanism is at work in
the Arctic. Thus, good models can be very useful in studying new data.
460
There are two models favored by most scientists in this area. Some scien-
tists put forth a chemical model that says the depletion is caused by chem-
ical events promoted by the presence of chlorofluorocarbons created by
industrial processes. Acceptance of this model was promoted by the dis-
covery of fluorine in the stratosphere. Fluorine does not naturally occur
there, but it is related to CFCs. The other model assumes that the ozone
hole was formed by dynamic air movement and mixing. This model best
fits data gathered by ozone-sensing balloons that sample altitudes up to 30
kilometers and then radio the data back to Earth. Ozone depletion is con-
fined to air between 12 and 20 kilometers. While the total ozone depletion
is 35 percent, different strata showed various amounts of depletion from 70
to 90 percent. Surprisingly, about half the ozone was gone in twenty-five
days. This finding does not fit the chemical model very well.
Besides ozone-sensing balloons, satellites are of much help. The Na-
tional Aeronautics and Space Administration (NASA) obtains measure-
ments with its Nimbus 7 satellite. Ozone measurements made by this satel-
lite helped to develop flight plans for the specialized aircraft NASA also
deploys in ozone studies. NASA’s ER-2 aircraft is a modified U-2 recon-
naissance plane that carries instruments up to 20 kilometers in altitude for
seven-hour flights to 80 degrees north latitude. A DC-8, operating during
the same period, is able to survey the polar vortex, owing to its greater
range. In addition, scientists utilize many meteorological techniques and
instruments, including chemical analysis of gases by means of infrared
spectroscopy, mass spectroscopy and gas spectroscopy combined, gas
chromatography, and oceanographic analysis of planktonic life in the
southern Atlantic, Pacific, and Indian oceans. As new research methods
become available, they are applied to this essential study.
Public Health Concerns

Atmospheric ozone provides a gauze of protection from the lethal effects
of ultraviolet radiation from the sun. This ability to absorb ultraviolet radi-
ation protects all life-forms on the earth’s surface from excessive ultravio-
let radiation, which destroys the life of plant and animal cells. Currently,
between 10 and 30 percent of the sun’s ultraviolet B (UV-B) radiation
reaches the earth’s surface. If ozone levels were to drop by 10 percent, the
amount of UV-B radiation reaching Earth would increase by 20 percent.
Present-day UV-B levels are responsible for the fading of paints and the
yellowing of window glazing and for car finishes becoming chalky. These
kinds of degradation will accelerate as the ozone layer is depleted. There
could also be increased smog, urban air pollution, and a worsening of the
problem of acid rain in cities. In humans, UV-B causes sunburn, snow
461
blindness, skin cancer, cataracts, and excessive aging and wrinkling of

skin. Skin cancer is the most common form of cancer—more than 400,000
new cases are reported every year in the United States alone. The National
Academy of Sciences has estimated that each 1 percent decline in ozone
would increase the incidence of skin cancer by 2 percent. Therefore, a 3 per-
cent depletion in ozone would produce some 20,000 more cases of skin
cancer in the United States every year.
Ecological Concerns
Many other forms of life—from bacteria to forests and crops—are ad-
versely affected by excessive radiation as well. Ultraviolet radiation affects
plant growth by slowing photosynthesis and by delaying germination in
many plants, including trees and crops. Scientists have a great concern for
the organisms that live in the ocean and the effect ozone depletion may
have on them. Phytoplankton, zooplankton, and krill (a shrimplike crusta-
cean) could be greatly depleted if there were a drastic increase in ultravio-
let A and B. The result would be a tremendous drop in the population of
these free-floating organisms. These organisms are important because they
are the beginning of the food chain. Phytoplankton use the energy of sun-
light to convert inorganic compounds, such as phosphates, nitrates, and
silicates, into organic plant matter. This process provides food for the next
step in the food chain, the herbivorous zooplankton and krill. They, in turn,
become the food for the next higher level of animals in the food chain. Ini-
tial studies of this food chain in the Antarctic suggest that elevated levels of
ultraviolet radiation impair photosynthetic activity. Recent studies show
that a fifteen-day exposure to UV-B levels 20 percent higher than normal
can kill off all anchovy larvae down to a depth of 10 meters. There is also
concern that ozone depletion may alter the food chain and even cause
changes in the organism’s genetic makeup. An increase in the ultraviolet
radiation is likely to lower fish catches and upset marine ecology, which
has already suffered damage from human-made pollution. On a world-
wide basis, fish presently provides 18 percent of all the animal protein con-
sumed.
International Response
The United Nations Environmental Program (UNEP) is working with gov-
ernments, international organizations, and industry to develop a frame-
work within which the international community can make decisions to
minimize atmospheric changes and the effects they could have on the
earth. In 1977, UNEP convened a meeting of experts to draft the World
Plan of Action on the Ozone Layer. The plan called for a program of re-
462
search on the ozone layer and on what would happen if the layer were
damaged. In addition, UNEP created a group of experts and government
representatives who framed the Convention for the Protection of the
Ozone Layer. This convention was adopted in Vienna in March, 1985, by
twenty-one states and the European Economic Community and has subse-
quently been signed by many more states. The convention pledges states
that sign to protect human health and the environment from the effects of
ozone depletion. Action has already been taken to protect the ozone layer.
Several countries have restricted the use of CFCs or the amounts pro-
duced. The United States banned the use of CFCs in aerosols in 1978. Some
countries, such as Belgium and the Nordic countries, in effect banned CFC
production altogether.
George K. Attwood and Jeffrey A. Joens
See also: Acid deposition; Deforestation; Global warming; Greenhouse ef-

fect; Pollution effects; Rain forests and the atmosphere.

Bast, Joseph L., Peter J. Hill, and Richard C. Rue. Eco-Sanity: A Common
Sense Guide to Environmentalism. Lanham, Md.: Heartland Institute,
1994.
Cagin, Seth, and Philip Dray. Between Earth and Sky: How CFCs Changed Our
World and Endangered the Ozone Layer. New York: Pantheon Books, 1993.
Firor, John. The Changing Atmosphere: A Global Challenge. New Haven,
Fisher, David E. Fire and Ice: The Greenhouse Effect, Ozone Depletion, and Nu-
clear Winter. New York: Harper & Row, 1990.
Graedel, T. E., and Paul J. Crutzen. Atmospheric Change: An Earth System
Perspective. New York: W. H. Freeman, 1993.
Roan, Susan. Ozone Crisis: The Fifteen-Year Evolution of a Sudden Global Emer-
gency. New York: John Wiley & Sons, 1989.
Rowland, F. Sherwood. “Stratospheric Ozone Depletion.” Annual Review of
Physical Chemistry 42 (1991): 731.
Shell, E. R. “Weather Versus Chemicals.” The Atlantic 259 (May, 1987): 27-
31.
Somerville, Richard C. J. The Forgiving Air: Understanding Environmental
Change. Los Angeles: University of California Press, 1996.
463
PALEOECOLOGY
Paleoecology is the study of ancient organisms and their relationships to one an-
other and to their environments. The characteristics of ancient environments may
be determined by examining rock and fossil features.
A s a field of science, paleoecology is most closely related to paleontol-

ogy, the study of fossils. It is also related to paleoclimatology, paleo-
geography, and a number of other areas of study dealing with the distant
past. All these disciplines have a handicap in common: Because they deal
with the past, scientists are unable to apply the usual scientific criteria of
direct observation and measurement of phenomena. Therefore, they look
to a number of different methods whereby evidence of past conditions and
organisms can be deduced.
Dendrochronology
One of the most intensively investigated paleoecological problems has been
the changing environments associated with the ice ages of the past million
years. Analysis of pollen from bogs in many parts of the world indicates that
there have been at least four advances and retreats of glaciers during that
period. Evidence for this is the changing proportions of pollen from tree
species found at the various depths of bogs. In North America, for exam-
ple, spruces (indicators of cool climate) formerly lived much farther south
than they do now. They were largely replaced almost eight thousand years
ago by other tree species, such as oaks, which are indicative of warmer cli-
mates. This warming trend was a result of the latest glacial retreat.
Tree-ring analysis, also known as dendrochronology, not only enables
paleoecologists to date past events such as forest fires and droughts but
also allows them to study longer-term cycles of weather and climate, espe-
cially those of precipitation and temperature. In addition, trees serve as ac-
cumulators of past mineral levels in the atmosphere and soil. Lead levels of
tree wood showed a sharp increase as the automobile became common in
the first half of the twentieth century because of lead additives in gasoline.
Tree rings formed since the 1970’s have shown a decrease in lead because
of the decline in use of leaded fuels. Tree-ring analysis has also been a valu-
able tool for archaeologists’ study of climatic changes responsible for shift-
ing patterns of population and agriculture among native Americans of the
southwestern United States.
464
Paleoecology
The Fossil Record

Fossil evidence is the chief source of paleoecological information. A fossil
bed of intact clam shells with both valves (halves) present in most individ-
uals, for example, usually indicates that the clams were preserved in the
site in which they lived (called autochthonous deposition). Had they been
transported by currents or tides to another site of deposition (allochthonous
deposition), the valves would have been separated, broken, and worn.
Similarly, many coal beds have yielded plant fossils that indicate that their
ancient environments were low-lying swamp forests with sluggish drain-
age periodically flooded by water carrying a heavy load of sand. The re-
sulting fossils may include buried tree stumps and trunks with roots still
embedded in their original substrate and numerous fragments of twigs,
leaves, and bark within the sediment.
Certain dome- or mushroom-shaped structures called stromatolites are
found in some of the most ancient of earth’s sedimentary rocks. These
structures may be several meters in diameter and consist of layers of mate-
rial trapped by blue-green algae (cyanobacteria). Such structures are cur-
rently being formed in shallow, warm waters. Uniformitarian interpreta-
tion of the three-billion-year-old stromatolites is that they were formed
under similar conditions. Their frequent association with mud cracks and
other shallow- and above-water features leads to the interpretation that
they were formed in shallow inshore environments subject to frequent ex-
posure to the air.
Relative oceanic temperature can be estimated by observing the direc-
tion in which the shells of certain planktonic organisms coil. The shell of
Globigerina pachyderma coils to the left in cool water and to the right in
warmer water. Globigerina menardii shells coil in an opposite fashion—to
the right in cool water and to the left in warmer water. Uniformitarian the-
ory leads one to believe that ancient Globigerina populations responded to
water temperature in a similar manner. Sea-bottom core samples showing
fossils with left- or right-coiling shells may be used to determine the rela-
tive water temperature at certain periods. Eighteen-thousand-year-old
sediments taken from the Atlantic Ocean show a high frequency of left-
handed pachyderma and right-handed menardii shells. Such observations
indicate that colder water was much farther south about eighteen thou-
sand years ago, a date that corresponds to the maximum development of
the last Ice Age.
Fossil Deposition
Fossil arrangement and position can be a clue to the environments in
which the organisms lived or in which they were preserved. Sea-floor cur-
465
Paleoecology
rents can align objects such as small fish and shells. Not only can the exis-
tence of the current be inferred, but also its direction and velocity can be
determined. Currents and tides can create other features in sediments
which are sometimes indicators of environment. If a mixture of gravel,
sand, silt, and clay is being transported by a moving body of water such as
a stream, tide, or current, the sediments will often become sorted by the
current and be deposited as conglomerates—sandstones, siltstones, and
shales. Such graded bedding can be used to determine the direction and
velocity of currents. Larger particles, such as gravel, would tend to be de-
posited nearer the sediment source than smaller particles such as clay. Sim-
ilarly, preserved ripple marks indicate current direction. Mud cracks in a
rock layer indicate that the original muddy sediment was exposed to the
atmosphere at least for a time after its deposition.
Fossil Composition
Certain minerals within fossil beds or within the fossil remains themselves
can sometimes be used to interpret the paleoenvironment. The presence of
pyrite in a sediment almost always indicates that the sedimentary environ-
ment was deficient in oxygen, and this, in turn, often indicates deep, still
water. Such conditions exist today in the Black Sea and even in some deep
lakes, with great accumulations of dead organic matter.
The method of preservation of the remains of the fossilized organism can
be an indication of the environment in which the creature lived (or died).
Amber, a fossilized resin, frequently contains the embedded bodies of
ancient insects trapped in the resin like flies on flypaper. This ancient envi-
ronment probably contained resin-bearing plants (mostly conifers), and
broken limbs and stumps that oozed resin to trap these insects. Mum-
mified remains in desert areas and frozen carcasses in the northern tundra
indicate the environments in which the remains were preserved thousands
of years ago.
Marks made on fossil parts by other organisms offer indirect evidence
of the presence and activity of other species that might not have left fossil
remains. Predators and scavengers can leave such marks on bones and
shells by boring, scratching, and gnawing. One of the most controversial
taphonomic problems in paleoecology is distinguishing between tooth
marks left by animal scavengers and predators on bones and those marks
left by the stone and bone tools of early human ancestors.
Fossil Assemblages and Trace Fossils

Fossil assemblages (thanatocoenoses) are the most commonly used indica-
tors of ancient environments. The use of any fossil in interpreting the past
466
Paleoecology
must be subject to several qualifications. The fossil record is sparse for

most groups of organisms because fossilization itself is a relatively rare
event. Rapid burial of the remains and the presence of hard body parts
(wood, shells, bones, and teeth) are only two of several fossilization pre-
requisites that must usually be met. This means that terrestrial organisms
and soft-bodied organisms are seldom fossilized. Events leading to fossil-
ization after the death of an organism (taphonomy) usually destroy the soft
tissues through decay and scavenging and often disrupt and distort the re-
maining hard parts through transportation and weathering.
An additional taphonomic problem is encountered when clumps or
clusters of fossil remains are located. Without careful study, it is difficult
to determine whether these assemblages are truly representative of the
groupings of the organisms in life or if they are simply coincidental aggre-
gations of such items as shells and limbs that were swept together by cur-
rents or wind and thus not indicative of the living situation and environ-
ment. Because of limitations on the interpretation of ancient environments
by the use of fossilized body parts, trace fossils are often more reliable indi-
cators of environmental conditions. Trace fossils are preserved tracks, bur-
rows, trails, and other indirect indications of the presence of an organism.
The presence of marine worm burrows, for example, can indicate environ-
mental factors such as salinity and depth. Such traces are not transported
from one site to another. Transportation results in their destruction. When-
ever these imprints are found, therefore, paleoecologists are able to make
some inferences about the environment in which they were formed.
Stratigraphy
One of the most important methods to be mastered by paleoecologists is
stratigraphy, the science of correlating and determining the age of rock lay-
ers with those of the fossils contained within these layers or formations.
Rock layers or strata are not usually connected over large regions. While
they might have been deposited as sediments at the same time and under
the same conditions, subsequent erosion has usually made the layers dis-
continuous. Stratigraphers attempt to correlate discontinuous rock strata
by measuring and describing them and by noting the presence of unique
fossils called index fossils. If two strata are correlated, then they were prob-
ably deposited during approximately the same period, although there may
be a gradation of conditions.
For example, there may be a layer of sediment deposited at the same
time, but under nearshore conditions at one spot and under offshore con-
ditions at another. Relative ages are determined by using the law of super-
position: Older rocks lie beneath younger rocks. One can say that a certain
467
Paleoecology
stratum is older than, the same age as, or younger than another layer, de-
pending upon their relative positions. Absolute ages (estimated age in
years before the present) are determined by measuring the amounts of cer-
tain radioactive elements within igneous rocks. Such radiometric age de-
terminations are of less value for sedimentary rocks since they give the age
of the minerals of the rock, not the age of the rock itself.
Related Fields
Paleoecological data are applicable to other, related paleo-fields of the
earth and life sciences. The study of fossils, paleontology, is enhanced by
the inclusion of information about the fossil organisms’ environments and
relationships with other organisms. Paleontologists should attempt to re-
construct ancient environments because organisms did not exist alone or in
vacuums: They lived in dynamic biological communities. Paleogeography
relies heavily on paleoecological information to discern the locations, di-
rections, and time intervals of glaciation, deposition of sediments, temper-
ature, and other environmental variables. This information has been used
to determine the past positions of continents and has been a valuable con-
tribution to scientists’ knowledge of continental drift.
Paleoclimatologists, who study ancient regional and planetwide condi-
tions, must make use of local bits of paleoecological information to see the
big picture of climate. One of the major concerns of paleoclimatology is the
recognition of planetary climatic cycles and associated environmental and
biological cycles. If there is a repeated recurrence of global environmental
change, then predictions about future climatic change become more accu-
rate and probable.
P. E. Bostick

chronology; Evolution: definition and theories; Evolution: history; Evo-
lution of plants and climates; Extinctions and evolutionary explosions;
Natural selection; Nonrandom mating, genetic drift, and mutation; Punc-
tuated equilibrium vs. gradualism; Speciation; Species loss.

Agashe, Shripad N. Paleobotany: Plants of the Past, Their Evolution, Paleo-
environment, and Application in Exploration of Fossil Fuels. Enfield, N.H.:
Science Publishers, 1997.
Arduini, Paolo, and Giorgio Teruzzi. Simon and Schuster’s Guide to Fossils.
New York: Simon & Schuster, 1986.
468
Paleoecology
Bennett, K. D. Evolution and Ecology: The Pace of Life. New York: Cambridge
Brett, Carlton E., and Gordon C. Baird, eds. Paleontological Events: Strati-
graphic, Ecological, and Evolutionary Implications. New York: Columbia
Cowen, Richard. History of Life. 3d ed. Malden, Md.: Blackwell Science,
2000.
Davis, Richard A. Depositional Systems: A Genetic Approach to Sedimentary
Geology. Englewood Cliffs, N.J.: Prentice-Hall, 1983.
Dodd, J. Robert, and Robert J. Stanton, Jr. Paleoecology: Concepts and Applica-
tions. 2d ed. New York: John Wiley & Sons, 1990.
National Research Council. Commission on Geosciences, Environment,
and Resources. Board on Earth Sciences and Resources. Effects of Past
Global Change on Life. Washington, D.C.: National Academy Press, 1995.
Newton, Cathryn, and Léo Laporte. Ancient Environments. 3d ed. Engle-
wood Cliffs, N.J.: Prentice-Hall, 1989.
Shipman, Pat. Life History of a Fossil: An Introduction to Taphonomy and Paleo-
ecology. Cambridge, Mass.: Harvard University Press, 1993.
469
PESTICIDES
Pesticides are substances designed to kill unwanted plants, fungi, or animals that
interfere, directly or indirectly, with human activities. The unintended impacts of
pesticides such as DDT have been to change ecosystems and their components.
T he major types of pesticides in common use are insecticides (to kill in-
sects), nematocides (to kill nematodes), fungicides (to kill fungi), her-
bicides (to kill weeds), and rodenticides (to kill rodents). Herbicides and
insecticides make up the majority of the pesticides applied in the environ-
ment. Biopesticides are beneficial microbes, fungi, insects, or animals that
kill pests. While the use of pesticides has mushroomed since the introduc-
tion of monoculture (the agricultural practice of growing only one crop on
a large amount of acreage), the application of toxins to control pests is by
no means new.
Insecticides: History of Use

The use of sulfur as an insecticide dates back before 500 b.c.e. Salts from
heavy metals such as arsenic, lead, and mercury were used as insecticides
from the fifteenth century until the early part of the twentieth century, and
residues of these toxic compounds are still being accumulated in plants
that are grown in soil where these materials were used. In the seventeenth
and eighteenth centuries, natural plant extracts, such as nicotine sulfate
from tobacco leaves and rotenone from tropical legumes, were used as in-
secticides. Other natural products, such as pyrethrum from the chrysan-
themum flower, garlic oil, lemon oil, and red pepper, have long been used
to control insects.
In 1939 the discovery of dichloro-diphenyl-trichlorethane (DDT) as a
strong insecticide opened the door for the synthesis of a wide array of syn-
thetic organic compounds to be used as pesticides. Chlorinated hydrocar-
bons such as DDT were the first group of synthetic pesticides. Other com-
monly used chlorinated hydrocarbons have in the past included aldrin,
endrin, lindane, chlordane, and mirex. Because of their low biodegrad-
ability and persistence in the environment, they proliferated up the food
chain and became concentrated in predators, such as birds and animals
butchered for human consumption. The use of these compounds was
therefore banned or severely restricted in the United States, but only after
years of use.
470
Pesticides
Organophosphates such as malathion, parathion, and methamidophos

have replaced the chlorinated hydrocarbons. These compounds biode-
grade in a fairly short time but are generally much more toxic to humans
and other animals than the compounds they replaced. In addition, they are
water-soluble and therefore more likely to contaminate water supplies.
Carbamates such as carbaryl, maneb, and aldicarb have also been used in
place of chlorinated hydrocarbons. These compounds rapidly biodegrade
and are less toxic to humans than organophosphates, but they are less ef-
fective in killing insects.
Herbicides
Herbicides are classified according to their method of killing rather than
their chemical composition. As their name suggests, contact herbicides
such as atrazine and paraquat kill when they come in contact with a plant’s
leaf surface. Contact herbicides generally disrupt the photosynthetic mecha-
nism. Systemic herbicides such as diuron and fenuron circulate through-
out the plant after being absorbed. They generally mimic the plant hor-
mones and cause abnormal growth to the extent that the plant can no
longer supply sufficient nutrients to support growth. Soil sterilants such as
triflurain, diphenamid, and daiapon kill microorganisms necessary for
plant growth and also act as systemic herbicides.
The spraying of herbicides and pesticides remains a standard agricultural practice

in the cultivation of both food crops and ornamental crops such as these tulips.
(PhotoDisc)
471
Pesticides
Current Use
In the United States, approximately 55,000 different pesticide formulations
are available, and Americans apply about 500 million kilograms (1.1 bil-
lion pounds) of pesticides each year. Fungicides account for 12 percent of
all pesticides used by farmers, insecticides account for 19 percent, and her-
bicides account for 69 percent. These pesticides have been used primarily
on four crops: soybeans, wheat, cotton, and corn. Approximately $5 billion
is spent each year on pesticides in the United States, and about 20 percent
of this is for nonfarm use. On a per-unit-of-land basis, homeowners apply
approximately five times as much pesticide as do farmers. On a worldwide
basis, approximately 2.5 tons (2,270 kilograms) of pesticides are applied
each year. Most of these chemicals are applied in developed countries, but
the amount of pesticide used in developing countries is rapidly increasing.
Approximately $20 billion is spent worldwide each year, and this expendi-
ture is expected to increase in the future, particularly in the developing
countries.
Despite current concerns about their toxicity and biomagnification,
pesticide use has had a beneficial impact on the lives of humans by in-
creasing food production and reducing food costs. Even with pesticides,
pests reduce the world’s potential food supply by as much as 55 percent.
Without pesticides, this loss would be much higher, resulting in increased
starvation and higher food costs. Pesticides also increase the profit mar-
gin for farmers. It has been estimated that for every dollar spent on pes-
ticides, farmers experience an increase in yield worth three to five dol-
lars. Pesticides appear to work better and faster than alternative methods
of controlling pests. These chemicals can rapidly control most pests, are
cost-effective, can be easily shipped and applied, and have a long shelf
life compared to alternative methods. In addition, farmers can quickly
switch to another pesticide if genetic resistance to a given pesticide de-
velops.
Perhaps the most compelling argument for the use of pesticides is
the fact that pesticides have saved lives. It has been suggested that since
the introduction of DDT, the use of pesticides has prevented approxi-
mately seven million premature human deaths from insect-transmitted
diseases such as sleeping sickness, bubonic plague, typhus, and malaria.
Perhaps even more lives have been saved from starvation because of
the increased food production resulting from the use of pesticides. It has
been argued, therefore, that this one benefit outweighs the potential health
risks of pesticides. In addition, new pesticides are continually being devel-
oped, and safer and more effective pest control may be available in the
future.
472
Pesticides
The publication of Rachel Carson’s

classic Silent Spring (1962), which
outlined the ecotoxicity of the pesticide
DDT, helped stimulate the
environmental movement of the 1960’s.
(Library of Congress)
Environmental Concerns
In spite of all the advantages of using pesticides, their benefit must be bal-
anced against the potential environmental damage they can cause. An
ideal pesticide would have the following characteristics: It should not kill
any organism other than the target pest; it would in no way affect the
health of nontarget organisms; it would degrade into nontoxic chemicals
in a relatively short time; it would prevent the development of resistance in
the organism it is designed to kill; and it would be cost-effective. Since no
currently available pesticide meets all of these criteria, a number of envi-
ronmental problems have developed, one of which is broad-spectrum poi-
soning. Most, if not all, chemical pesticides are not selective; they kill a
wide range of organisms rather than just the target pest. Killing beneficial
insects, such as bees, lady bird beetles, and wasps, may result in a range of
problems. For example, reduced pollination and explosions in the popula-
tions of unaffected insects can occur.
When DDT was first used as an insecticide, many people believed that it
was the final solution for controlling many insect pests. Initially, DDT dra-
matically reduced the number of problem insects; within a few years, how-
ever, a number of species had developed genetic resistance to the chemical
and could no longer be controlled with it. By the 1990’s there were approxi-
mately two hundred insect species with genetic resistance to DDT. Other
chemicals were designed to replace DDT, but many insects also developed
resistance to these newer insecticides. As a result, although many synthetic
473
Pesticides
chemicals have been introduced to the environment, the pest problem is

still as great as it ever was.
Depending on the type of chemical used, pesticides remain in the envi-
ronment for varying lengths of time. Chlorinated hydrocarbons, for exam-
ple, can persist in the environment for up to fifteen years. From an eco-
nomic standpoint, this can be beneficial because the pesticide has to be
applied less frequently, but from an environmental standpoint, it is detri-
mental. In addition, when many pesticides are degraded, their breakdown
products, which may also persist in the environment for long periods of
time, can be toxic to other organisms.
Pesticides may concentrate as they move up the food chain, a process
called biomagnification. All organisms are integral components of at least
one food pyramid. While a given pesticide may not be toxic to species at the
base, it may have detrimental effects on organisms that feed at the apex be-
cause the concentration increases at each higher level of the pyramid. With
DDT, for example, some birds can be sprayed with the chemical without any
apparent effect, but if these same birds eat fish that have eaten insects that
contain DDT, they lose the ability to metabolize calcium properly. As a re-
sult, they lay soft-shelled eggs, which causes deaths of most of the offspring.
Pesticides can be hazardous to human health. Many pesticides, particu-
larly insecticides, are toxic to humans, and thousands of people have been
killed by direct exposure to high concentrations of these chemicals. Many
of these deaths have been children who were accidentally exposed to toxic
pesticides because of careless packaging or storage. Numerous agricul-
tural laborers, particularly in developing countries where there are no
stringent guidelines for handling pesticides, have also been killed as a re-
sult of direct exposure to these chemicals. Workers in pesticide factories are
also a high-risk group, and many of them have been poisoned through job-
related contact with the chemicals. Pesticides have been suspected of caus-
ing long-term health problems such as cancer. Some of the pesticides have
been shown to cause cancer in laboratory animals, but there is currently no
direct evidence to show a cause-and-effect relationship between pesticides
and cancer in humans.
D. R. Gossett
See also: Biomagnification; Biopesticides; Genetically modified foods; In-

tegrated pest management; Soil contamination.

Altieri, Miguel A. Agroecology: The Science of Sustainable Agriculture. 2d ed.
Boulder, Colo.: Westview Press, 1995.
474
Pesticides
Carson, Rachel. Silent Spring. 1962. Reprint. Thorndike, Maine: G. K. Hall,

1997.
Mannion, Antionette M., and Sophia R. Bowlby. Environmental Issues in the
1990’s. Chichester, N.Y.: John Wiley & Sons, 1992.
Milne, George W. A., ed. Ashgate Handbook of Pesticides and Agricultural
Chemicals. Burlington, Vt.: Ashgate, 2000.
Nadakavukaren, Anne. Man and Environment: A Health Perspective. 2d ed.
Prospect Heights, Ill.: Waveland Press, 1990.
Pierce, Christine, and Donald VanDeVeer. People, Penguins, and Plastic
Trees. 2d ed. Belmont, Calif.: Wadsworth, 1995.
475
PHEROMONES
ecology
Pheromones are chemicals or mixtures of chemicals that are used as messages be-
tween members of a species. They are integral parts of the social communication
within most species. They may prove to be of great value in pest control and in en-
hancing agricultural production.
P heromones are chemical signals. Originally defined to include only

signals between individuals of the same animal species, the term has
been generalized to designate any chemical or chemical mixture that,
when released by one member of any species, affects the physiology or be-
havior of another member of the same species. “Pheromone” is also one of
a set of terms developed to express the chemical interactions in ecological
communities.
All pheromones are semiochemicals that carry information between
members of a single species. To do so, the pheromone must be released into
the atmosphere or placed on some structure in the organism’s environ-
ment. It is thus made available to other members of the species for interpre-
tation and response. It is also available to members of other species, how-
ever, so it is a potential allelochemical.
Types of Pheromones
There are two general types of pheromone: those that elicit an immediate
and predictable behavioral response, called releaser or signal pheromones,
and those that bring about a less obvious physiological response, called
primer pheromones (because they prime the system for a possible behav-
ioral response). Pheromones are also categorized according to the mes-
sages they carry. There are trail, marker, aggregation, attractant, repellant,
arrestant, deterrent, stimulant, alarm, and other pheromones. Their func-
tions are suggested by the terms used to name them.
To appreciate fully the complexity of the interactions under consider-
ation, it is important to remember that a pheromone may also be acting as a
kairomone, allomone, or hormone. For example, klipspringer antelope
mark vegetation in their environment with a chemical secreted from a spe-
cial gland. Other klipspringer investigate the marks to gather information
on the marking individual. Ticks that parasitize the klipspringer, however,
are also attracted to the chemical marks and thus increase their chance of
476
Pheromones
attaching to a host when the mark is renewed or when another klipspringer

investigates it. The tick is using the pheromone as a kairomone. Phero-
mones can act as allomones as well, though the interaction is sometimes
less direct. Bolas spiders produce the sex-attractant pheromone of a female
moth and use it to lure male moths to a trap. The spider uses the moth
pheromone as an allomone.
Pheromonal Compounds and Strategies

The chemical compounds that act as pheromones are numerous and di-
verse. Most are lipids or chemical relatives of the lipids, including many
steroids. Even a single pheromonal message may require a number of dif-
ferent compounds, each present in the proper proportion, so that the active
pheromone is actually a mixture of chemical compounds.
Different physical and chemical characteristics are required for phero-
mones with different functions. Attractant pheromones must generally be
volatile to permit atmospheric dispersal to their targets. Many female in-
sects emit sex-attractant pheromones to advertise their readiness to mate.
The more widely these can be dispersed, the more males the advertisement
will reach. On the other hand, many marking pheromones need not be es-
pecially volatile because they are placed at stations which are checked pe-
riodically by the target individuals. The klipspringer marking pheromone
is an example. Some pheromones are exchanged by direct contact, and
these need not have any appreciable volatile component. Many mammals
rub, lick, and otherwise contact one another in social contexts and ex-
change pheromones at these times.
Specificity also varies for pheromones with different functions. Sex at-
tractants usually need to be very specific, directed only to members of the
opposite sex and the same species. Alarm pheromones, on the other hand,
need not be so specific. These pheromones simply alert other members
of the same species to a disturbance. It is usually harmless, and some-
times even helpful, to alert members of other species as well. In keeping
with this argument, related groups of ant species produce species-specific
sex-attractant pheromones: Each female attracts only males of its own spe-
cies. In contrast, alarm pheromones of any species in the group will stimu-
late defensive reactions in individuals of many in the species.
Pheromonal systems are not organized in any standard way in different
species. Many mites and ticks also have nonspecific alarm pheromones.
Surprisingly, some groups also have nonspecific sex-attractant pheromones.
In these cases, the specificity necessary for reproductive efficiency is gener-
ated by species-specific mating stimulant pheromones. These pheromones
are produced by a female after males have been attracted to her. They stim-
477
Pheromones
ulate mating behavior, but only in males of the same species. Thus the re-
quired specificity is achieved by a different mechanism. This is only one of
many examples of the diversity of pheromonal schemes among organisms.
Pheromone Sources and Receptors

The sources of pheromones are also diverse. Some pheromones are pro-
duced by specialized glands; many insect species have glands specialized
for the production of pheromones. One example is the harvester ant’s
alarm pheromone, which is produced in the mandibular gland at the base
of the jaws. Other pheromones seem to be by-products of other bodily
functions. The lipids of mammalian skin are probably primarily important
in waterproofing and in maintaining the outer layer of the skin, but many
also function as pheromones.
The reproductive tract is an important source of pheromones in many
species. These usually act as sex-attractant or sex-stimulant pheromones or
as signal pheromones that give information on the sexual state of the emit-
ter. The urine and feces of many species also contain pheromones that are
used to mark territory boundaries and to transmit other information about
the marking individual. Many pheromones seem to be produced not by
the sending organism alone but by microorganisms living on the skin or in
the glands or cavities of the sender’s body. These microbes convert prod-
ucts of their host into the actual signal molecules, or pheromones, used by
the host.
The receptors for pheromones are also of many different types, and the
chemical receptors for taste and smell are often involved. In vertebrates,
the vomeronasal organ (Jacobson’s organ) seems to be an important recep-
tor for many pheromones. It is a pouch off the mouth or nasal passages,
and it contains receptors similar to those for smell. It is nonfunctional in
humans, but it functions in more primitive mammals and seems to be of
great importance to snakes and other reptiles. Insects and other inverte-
brates have many specialized structures for receiving pheromonal mes-
sages. Perhaps the best-known example is the feathery antennae of many
male moths, which are receptors for the female moth’s sex-attractant
pheromone. Some pheromones seem to be absorbed through the skin or in-
ternal body linings and to bring about their effects by attaching to some
unknown internal receptor.
Prevalence in Nature
Pheromones are widespread in nature, occurring in most, if not all, species.
Most are poorly understood. The best-known are those found in insects,
partly because of their potential use in the control of pest populations and
478
Pheromones
partly because the relative simplicity of insect behavior allowed for rapid
progress in the identification of pheromones and their actions. Despite
these advantages, much remains to be learned even about insect phero-
mones. Mammalian pheromones are not as well known, although they
may also be of economic importance. The more complex behavior of mam-
mals makes the study of their responses to pheromones much more diffi-
cult.
Research Methods
Both behavioral and chemical techniques are required to study phero-
mones and other semiochemicals. The observation of behavior, either in
nature or in captivity, often suggests pheromonal functions. These hypoth-
esized functions are then tested by presenting the pheromone to a poten-
tially responsive organism and observing the response. Situations may be
arranged which demand the subject’s response to a particular pheromone
under otherwise natural conditions. Alternatively, the organisms may be
observed in enclosures to help control the experimental context. The pre-
sentation of the hypothetical pheromone may be in the form of another or-
ganism of the same species or some structure to which the presumed
pheromone has been applied. The observed response (or lack of response)
gives information on the status of the presented chemical as a pheromone
in that behavioral context.
While the pheromonal function of secretions from a gland or other
source can be determined from these behavioral tests, the tests can give in-
formation on specific chemical compounds only if the compounds can be
isolated and identified. The isolation and identification of pheromonal
compounds are challenging because of the great complexity of the secre-
tions in which they are found and the exceptionally small amounts that are
required to elicit a response. Many separation and identification tech-
niques are used. One of the most powerful is a combination of gas chroma-
tography and mass spectrometry.
Gas chromatography is used to separate and sometimes to identify
chemicals that are volatile or can be made volatile. The unknown chemical
is mixed with an inert gas, called the mobile phase of the gas chromatogra-
phy system. This mixture is passed through a tube containing a solid,
called the stationary phase. The inert gas does not interact with the solid;
however, many of the compounds mixed with it do, each to an extent de-
termined by the characteristics of the compound and the characteristics of
the stationary phase. Some members of the mixture will interact very
strongly with the solid and so move slowly through the tube, whereas oth-
ers may not interact with the solid at all and so pass through rapidly. Other
479
Pheromones
members of the mixture interact at intermediate strengths and so spend in-

termediate amounts of time in the tube. The different compounds are re-
corded and collected separately as they exit from the tube.
For identification, the compounds are often passed on to a mass spec-
trometer. In mass spectrometry the compound is broken up into electri-
cally charged particles. The particles are then separated according to their
mass-to-charge ratio, and the relative number of particles of each mass-to-
charge ratio is recorded and plotted. The original compound can usually
be identified by the pattern produced under the specific conditions used.
After separation and identification, the individual chemicals may be sub-
jected to behavioral studies.
Uses for Pheromones

Pheromones and other semiochemicals are of interest simply from the
standpoint of understanding communication between living things. In ad-
dition, they have the potential to provide effective, safe agents for pest con-
trol. The possibilities include sex-attractant pheromones to draw pest in-
sects of a particular species to a trap (or to confuse the males and keep them
from finding females) and repellant pheromones to drive a species of in-
sect away from a valuable crop species. One reason for the enthusiasm
generated by pheromones in this role is their specificity. Whereas insecti-
cides generally kill valuable insects as well as pests, pheromones will often
be specific for one or a few species.
These chemicals were presented as a panacea for insect and other pest
problems in the 1970’s, but most actual attempts to control pest popula-
tions failed. Many people in the field have suggested that lack of under-
standing of the particular pest and its ecological context was the most com-
mon cause of failure. They maintain that pest-control applications must be
made with extensive knowledge and careful consideration of pest charac-
teristics and the ecological system. In this context, pheromones have be-
come a part of integrated pest management (IPM) strategies, in which they
are used along with the pest’s parasites and predators, resistant crop vari-
eties, insecticides, and other weapons to control pests. In this role, phero-
mones have shown great promise.
Some consideration has been given to the control of mammalian pests
with pheromones, though this field is not as well developed as that of in-
sect control. Pheromonal control of mammalian reproduction has received
considerable attention for other reasons: Domestic mammals are of great
economic importance, and many wild mammalian species are endangered
to the point that captive breeding has been attempted. The manipulation of
reproductive pheromones may be used to enhance reproductive potentials
480
Pheromones
in both cases. The complexity of mammalian behavioral and reproductive

systems, however, and the subtle changes brought about by mammalian
pheromones present a particular challenge. As with insect pest control, the
key to progress is a complete understanding of the entire system being ma-
nipulated.
Pheromones and other semiochemicals are of great potential economic
importance as substitutes for or adjuncts to toxic pesticides in pest man-
agement. Mammalian reproductive pheromones are being explored as
tools to enhance reproductive efficiency in domestic and endangered mam-
mals. A complete understanding of the complex roles of pheromones in
each of the systems being managed is necessary for success in all these en-
deavors.
Carl W. Hoagstrom
See also: Allelopathy; Biopesticides; Communication; Defense mecha-

nisms; Displays; Ethology; Insect societies; Isolating mechanisms; Metabo-
lites; Reproductive strategies.

Agosta, William C. Chemical Communication: The Language of Pheromones.
New York: Scientific American Library, 1992.
Albone, Eric S. Mammalian Semiochemistry: The Investigation of Chemical Sig-
nals Between Mammals. New York: John Wiley & Sons, 1984.
Booth, William. “Revenge of the ‘Nozzleheads.’ ” Science 239 (January 8,
1988): 135-137.
Carde, Ring T., and Albert K. Minks, eds. Insect Pheromone Research: New
Directions. New York: Chapman and Hall, 1997.
Mayer, Marion S., and John R. McLaughlin. Handbook of Insect Pheromones
and Sex Attractants. Boca Raton, Fla.: CRC Press, 1991.
Mitchell, Everett R., ed. Management of Insect Pests with Semiochemicals: Con-
cepts and Practice. New York: Plenum, 1981.
Mittler, Thomas E., Frank J. Radovsky, and Vincent H. Resh, eds. Annual
Review of Entomology. Vol. 46. Palo Alto, Calif.: Annual Reviews, 2000.
Nordlund, Donald A., Richard L. Jones, and W. Joe Lewis, eds. Semio-
chemicals: Their Role in Pest Control. New York: John Wiley & Sons, 1981.
Vandenbergh, John G., ed. Pheromones and Reproduction in Mammals. New
York: Academic Press, 1983.
Wilson, Edward O. “Pheromones.” Scientific American 206 (May, 1963): 100-
114.
481
PHYTOPLANKTON
Types of ecology: Ecoenergetics; Ecotoxicology
Most plankton are microscopic and are usually single-celled, a chain of cells, or a
loose group of cells. Algal and cyanobacterial plankton are referred to as phyto-
plankton.
T he term “plankton,” from Greek planktos for “wandering,” is applied

to any organism that floats or drifts with the movement of the ocean
water. Whereas the heterotrophic crustaceans and larvae of animals are re-
ferred to as zooplankton, phytoplankton phytoplankton (literally, “plant”
plankton) refer to a collection of diverse, largely algal and cyanobacterial,
microorganisms. The phytoplankton include diatoms, unicellular cyano-
bacteria and coccolithophorids in nutrient-poor waters, and cryptomo-
nads. They manufacture organic material from carbon dioxide, usually
through photosynthesis, and therefore occupy the key trophic level of pro-
ducers, at the base of the food chain. Phytoplankton are responsible for
one-half of the world’s primary photosynthesis and produce one-half of
the oxygen in the atmosphere.
Eighty to ninety percent of the weight of phytoplankton is water, with
the rest made up of protein, fat, salt, carbohydrates, and minerals. Some
species have compounds of calcium or silica that make up their shells or
skeletons. Phytoplankton include many of the algal phyla: Chrysophyta
(chrysophytes), Phaeophyta (golden-brown algae), coccolithophores, silico-
flagellates, and diatoms. The most common type of phytoplankton is the
diatom (phylum Bacillariophyta), a single-celled organism that can form
complex chains. Dinoflagellates (phylum Dinophyta) are the most complex
of the phytoplankton. They are unicellular and mobile. Green algae (phy-
lum Chlorophyta) are usually found in estuaries or lagoons in the late sum-
mer and fall. Some species can cause toxic algal blooms associated with
coastal pollution and eutrophication. Cyanobacteria (often called blue-
green algae but not true algae) are prominent near shore waters with lim-
ited circulation and brackish waters.
Role in the Food Chain

Phytoplankton are primary producers, responsible for half the world’s pri-
mary photosynthesis: the conversion of light energy and inorganic matter
into bioenergy and organic matter. Each year, 28 billion tons of carbon and
482
Phytoplankton
250 billion to 300 billion tons of photosynthetically produced materials are

generated in the oceans by phytoplankton. All animal organisms eliminate
carbon dioxide into the atmosphere, and plants remove carbon dioxide
from the air through photosynthesis. In the oceans’ carbon cycle, carbon
dioxide from the atmosphere dissolves in the ocean. Photosynthesis by
marine plants, mainly phytoplankton, converts the carbon dioxide into or-
ganic matter. Carbon dioxide is later released by plants and animals during
respiration, while carbon is also excreted as waste or in the dead bodies of
organisms. Bacteria decompose organic matter and release the carbon di-
oxide back into the water. Carbon may be deposited as calcium carbonate
in biogenous sediments and coral reefs (made of skeletons and shells of
marine organisms).
Because they are primary producers of organic matter through photo-
synthesis, phytoplankton play a key role in the world’s food chain: They
are its very beginning. Sunlight usually penetrates only 200 to 300 feet
deep into ocean waters, a region called the photic zone. Most marine plant
and animal life and feeding take place in this zone. Phytoplankton, the first
level in the marine food chain, are the primary food source for zooplank-
ton and larger organisms. These microscopic plants use the sun’s energy to
absorb minerals to make basic nutrients and are eaten by herbivores, or
plant eaters. Herbivores are a food source for carnivores, the meat eaters.
In temperate zones, phytoplankton increase greatly in the spring, decline
in the summer, and increase again in the fall. Zooplankton (animal plank-
ton) are at their maximum abundance after the spring increase, and their
grazing on the phytoplankton causes a decrease in phytoplankton popula-
tion in the summer. Fish and invertebrates that eat zooplankton become
more abundant and so on, up the food chain. Krill, planktonic crustaceans,
and larvae commonly eaten by whales, fish, seals, penguins, and seabirds
feed on diatom phytoplankton.
Red Tides
The term “red tide” is applied to red, orange, brown, or bright-green
phytoplankton blooms, or even to blooms that do not discolor the water.
Red tides are poorly understood and unpredictable. No one is certain what
causes the rapid growth of a single species of phytoplankton, although
they can blossom where sunlight, dissolved nutrient salts, and carbon
dioxide are available to trigger photosynthesis. Dense phytoplankton
blooms occur in stable water where lots of nutrients from sewage and run-
off are available. Natural events, such as storms and hurricanes, may
remobilize populations buried in the sediment. These nuisance blooms,
usually caused by dinoflagellates, which turn the water a reddish brown,
483
Phytoplankton
and cyanobacteria, are becoming more frequent in coastal waters, possibly

because of increased human populations and sewage. In shallower bodies
of water, such as bays and estuaries, nutrients from winter snow runoffs,
spring rains, tributaries, and sewage bring about spring and summer
blooms.
Some of the poisons produced during red tides are the most powerful
toxins known. The release of toxins by dinoflagellates may poison the
higher levels of the food chain as well as suppress other phytoplankton
species. These toxins cause high mortality in fish and other marine verte-
brates. They can kill the whales and seabirds that eat contaminated fish.
Dinoflagellates produce a deadly neurotoxin called saxitoxin, which is
fifty times more lethal than strychnine or curare. Commercial shellfish,
such as mussels, clams, and crabs, can store certain levels of the toxin in
their bodies.
People who eat contaminated shellfish may experience minor symp-
toms, such as nausea, diarrhea, and vomiting, or more severe symptoms
such as loss of balance, coordination and memory, tingling, numbness,
slurred speech, shooting pains, and paralysis. In severe cases, death results
from cardiac arrest. When the toxins are blown ashore in sea spray, they
can cause sore throats or eye and skin irritations.
Toxic blooms costs millions of dollars in economic losses, especially for
fisheries which cannot harvest some species of shellfish. Smaller fish farms
can be devastated. Additionally, coastal fish deaths foul beaches and shore
water with decaying bodies, which can cripple tourism in the coastal re-
gions.
Not all blooms are harmful, but they do affect the marine environment.
Even when no toxins are released, massive fish kills can result when the
large blooms of phytoplankton die. When the blooming phytoplankton
population crashes, bacterial decomposition depletes the oxygen in the
water, which in turn reduces water quality, and fish and other marine ani-
mals suffocate.
Virginia L. Hodges
See also: Biomass related to energy; Eutrophication; Food chains and

webs; Marine biomes; Nutrient cycles; Trophic levels and ecological
niches.

Castro, Peter, and Michael Huber. Marine Biology. 3d ed. Boston: McGraw-
Hill, 2000.
Cousteau, Jacques. The Ocean World. New York: Harry N. Abrams, 1985.
484
Phytoplankton
Levinton, Jeffrey S. Marine Biology: Function, Biodiversity, Ecology. New

Sumich, James L. An Introduction to the Biology of Marine Life. 5th ed.
Dubuque, Iowa: William C. Brown, 1992.
485
POISONOUS ANIMALS
ecology
Animal poisons, or venoms, are used both as a defense mechanism and as a preda-
tory strategy. These toxins can be delivered by biting, stinging, or body contact.
Poisonous species occur throughout the animal kingdom and include snakes, in-
sects, spiders and other arachnids, mammals, lizards, and fish.
S ubstances that cause disease symptoms, injure tissues, or disrupt life

processes on entering the body are poisons. When ingested in large
quantities, most poisons kill. Poisons can be contacted from minerals, in
vegetable foods, or through animal attack. Any poison of animal origin is a
venom. Venoms are delivered by biting, stinging, or other body contact.
These animal poisons are used to capture prey or in self-defense. Often, it
seems that the ability to make venom arose in animals that were too small,
too slow, or too weak to maintain an ecological niche otherwise.
The most familiar poisonous animals are snakes, insects, spiders, and
some other arachnids. Poisonous species, however, occur throughout the
animal kingdom, including a few mammals and lizards, and some fish. The
severity of venom effects depends on its chemical nature, the nature of the
contact mechanisms, the amount of venom delivered, and victim size. For ex-
ample, all spiders are poisonous. However, their venom is usually dispensed
in small amounts that do not affect humans. Hence, few spiders kill hu-
mans, though they kill prey and use venom in self-defense very effectively.
Chemically, venoms vary greatly. Snake venoms are mixtures of en-
zymes and toxins. Study of their effects led to the identification of hemo-
toxins, which cause blood vessel damage and hemorrhage; neurotoxins,
which paralyze nerves controlling heart action and respiration; and clot-
ting agents, which excessively promote or prevent blood clotting. Cobras,
coral snakes, and arachnids all have neurotoxic venoms.
Lizards, Arthropods, and Insects

Only two species of poisonous lizard are known: Gila monsters and beaded
lizards (both holoderms). They inhabit the southwestern deserts of the
United States and Mexico. They do not strike like snakes; rather, they bite,
hold on, and chew to apply their venom. Holoderm bites kill prey but
rarely kill humans. Beaded lizards grow to three feet long and Gila mon-
sters grow to two feet long.
486
Poisonous animals
Most poisonous arthropods are spiders and scorpions. Both use venom
to subdue or kill prey. As stated earlier, few spiders endanger humans be-
cause their venom is weak and is not injected in large quantities, but some
species have very potent venom and harm or even kill humans. Best
known of these are black widow spiders. Though rarely lethal to humans,
black widow bites cause cramps and paralysis.
All of the approximately six hundred scorpion species, of sizes between
one and ten inches, have tail-end stingers. Large, tropical scorpions can kill
humans, while American scorpions are smaller and less dangerous. Scor-
pions are more dangerous than spiders because they crawl into shoes and
other places where their habitat overlaps with that of humans.
Many insects, such as caterpillars, bees, wasps, hornets, and ants, use
venom in self-defense or to paralyze prey to feed themselves or offspring.
Caterpillars use poison spines for protection. Bees, wasps, hornets, and
ants use stingers for the same purpose. The venom of insects also kills
many organisms that seek to prey on them. Humans, however, are rarely
killed by insect bites. Such bites are usually mildly to severely painful for a
period from a few minutes to several days. However, for some humans
who are particularly sensitive, severe anaphylaxis occurs, in some cases
followed by death.
Poisonous Snakes
Poisonous snakes are colubrids, elapids, or vipers, depending on their ana-
tomic characteristics. All have paired, hollow fangs in the front upper jaw.
The fangs fold back against the upper palate when not used, and when a
snake strikes they swing forward to inject a venom that attacks the victim’s
blood and tissues. The heads of poisonous snakes are scale-covered and tri-
angular. Such snakes are found worldwide and include pit vipers, named
for the pits on each side of the head that contain heat receptors. The pits de-
tect warm-blooded prey, mostly rodents, in the dark. Pit vipers include rat-
tlesnakes, moccasins, copperheads, fer-de-lance, and bushmasters.
The populations and species of American and European poisonous
snakes differ. In North America, twenty such snake types occur: elapid
coral snakes and copperheads, sixteen rattler types, and cottonmouths (all
vipers). Vipers are found everywhere but Alaska. Rattlers have the widest
habitat, as shown by their abundance in the snake-rich Great Plains, Mis-
sissippi Valley, and southern Appalachia. In contrast, copperheads and
cottonmouths are abundant in Appalachia and the Mississippi Valley, re-
spectively. Mexican poisonous snakes are divided into two ranges: the
northern, from the U.S.-Mexican border to Mexico City, and the southern,
south of Mexico City. In the north, snakes are mostly rattlers, as in the con-
487
Poisonous animals
tiguous United States. Coral snakes and pit vipers are plentiful in the
south. Most perilous are the five- to eight-foot fer-de-lance, whose venom
kills many humans. All South American vipers live in tropical environ-
ments, except for rattlesnakes. Rattlers prefer arid environments, although
some are also found in tropical climates. Bushmasters, the largest South
American vipers, and elapid coral snakes are nocturnal and rarely endan-
ger humans. Tropical rattlers and lance-headed vipers, somewhat less noc-
turnal, kill many. Europe has few snakes, due to its cool climates and scarce
suitable habitats. Its few vipers range almost to the Arctic Circle. Eastern
Mediterranean regions hold most of the European vipers.
There are many poisonous snakes in Africa and Asia. North Africa,
mostly desert, has few snakes. Central Africa’s diverse poisonous snakes
are colubrid, elapid, and viper types. Elapids include dangerous black
mambas, twelve to fourteen feet long, and smaller cobras, which also occur
in South Africa. Among diverse vipers, the most perilous are Gaboon vi-
pers and puff adders. The Middle East, mostly desert, has few poisonous
snakes. Southeast Asia has the most poisonous snakes in the world, ela-
pids, colubrids, and vipers. This is due to snake habitats that range from
semiarid areas to rain forests. The huge human population explains why
this area has the world’s highest incidence of snakebite and related deaths.
Vipers bite most often, but elapids cause a larger portion of deaths. The Far
East snake population is complex, and its snakebite incidence is also high.
Its important poisonous snakes are pit vipers.
Australia and New Guinea have large numbers of poisonous snakes.
Australia has 65 percent of the world’s snakes, while New Guinea has 25
percent. Also, sea snakes occur offshore and in some rivers and lakes.
However, these countries have few snakebite deaths, because of the small
size and nocturnal nature of most of the indigenous snakes.
Poisonous Fish and Amphibians

Venomous fish are dangerous to those who enter the oceans, especially
fishermen who take them from their nets. The geographical distribution of
these fish is like all other fish. The highest population density is in warm
temperate or tropical waters. Numbers and varieties of poisonous fish de-
crease with proximity to the North and South Poles, and they are most
abundant in Indo-Pacific and West Indian waters.
A well-known group of poisonous fish, the stingrays (dasyatids), in-
habit warm, shallow, sandy-to-muddy ocean waters. Dasyatids lurk al-
most completely buried, awaiting prey that they sting to death with
barbed, venomous teeth in their tails. The tail poison is made in glands at
the bases of the teeth. Small, freshwater dasyatids are found in South
488
Poisonous animals
American rivers, such as the Amazon, hundreds of miles from the river
mouths. Stingrays near Australia grow to fifteen-foot lengths. The wide
distribution of stingrays and their danger to humans are mentioned in the
writings of Aristotle in the third century b.c.e. and they played a role in the
death of John Smith in 1608, who was killed by a stingray while exploring
Chesapeake Bay.
Also well known are the venomous Scorpaenidae fish family, many mem-
bers of which cause very painful stings. Zebrafish and stonefish are good ex-
amples. Both, like all scorpaenids, have sharp spines supporting dorsal fins.
The spines, used in self-defense, have venom glands. The most deadly fish
venom is that of the stonefish, which, when stepped on, can kill humans.
Frogs and Toads

Poisonous animals that endanger by contact are exemplified not only by
the zebrafish and stonefish just mentioned but also by poisonous frogs or
toads. Most such frogs and toads live in Africa and South America. Poison
dart frogs, for example, secrete poisons through the skin. In humans, the
effects of contact with these poisons range from severe irritation to death.
The poisons frighten away or kill most predators that attempt to eat the
frogs.
Ecological Significance
The ecological function of poisonous animals is to keep down the popula-
tion of insects, rodents, arachnids, and small fish. They thus contribute to
maintaining the balance of nature. Poisonous land animals, such as scorpi-
ons and many poisonous snakes, are often nocturnal and add another di-
mension to pest control by nighttime predation.
Sanford S. Singer

Metabolites; Pheromones; Poisonous plants; Predation.

Aaseng, Nathan. Poisonous Creatures. New York: Twenty-first Century
Books, 1997.
Edström, Anders. Venomous and Poisonous Animals. Malabar, Fla.: Krieger,
1992.
Foster, Steven, and Roger Caras. Venomous Animals and Poisonous Plants.
Boston: Houghton Mifflin, 1994.
Grice, Gordon D. The Red Hourglass: Lives of the Predators. New York:
Delacorte, 1998.
489
POISONOUS PLANTS
Types of ecology: Chemical ecology; Physiological ecology
Poisonous plants have evolved toxic substances that function to defend them
against herbivores and thereby better adapt them for survival.
A fter evolving adaptations that facilitated colonization of terrestrial

habitats, plants were confronted with a different type of problem.
This was the problem of herbivory, or the inclination of many different
types of organisms, from bacteria to insects to four-legged herbivores, to
eat plants. Pressures from herbivory drove many different types of plants,
from many different families, to evolve defenses. Some of these defenses
included changes in form, such as the evolution of thorns, spikes, or
thicker, tougher leaves. Other plants evolved to produce chemical com-
pounds that make them taste bad, interrupt the growth and life cycles of
the herbivores, make the herbivores sick, or kill them outright.
Phytochemicals
One of the most interesting aspects of plants, especially prevalent in the an-
giosperms (flowering plants), is their evolution of substances called sec-
ondary metabolites, sometimes referred to as phytochemicals. Once con-
sidered waste products, these substances include an array of chemical
compounds: alkaloids, quinones, essential oils, terpenoids, glycosides (in-
cluding cyanogenic, cardioactive, anthraquinone, coumarin, and sapo-
nin glycosides), flavonoids, raphides (also called oxalates, which contain
needle-like crystals of calcium oxalate), resins, and phytotoxins (highly
toxic protein molecules). The presence of many of these compounds can
characterize whole families, or even genera, of flowering plants.
Effects on Humans
The phytochemicals listed above have a wide range of effects. In humans,
some of these compounds will cause mild to severe skin irritation, or con-
tact dermatitis; others cause mild to severe gastric distress. Some cause hal-
lucinations or psychoactive symptoms. The ingestion of many other types
of phytochemicals proves fatal. Interestingly, many of these phytochem-
icals also have important medical uses. The effects of the phytochemicals
are dependent on dosage: At low doses, some phytochemicals are thera-
peutic; at higher doses, some can kill.
490
Poisonous plants
Alkaloids
Alkaloids are nitrogenous, bitter-tasting compounds of plant origin. More
than three thousand alkaloids have been identified from about four thou-
sand plant species. Their greatest effects are mainly on the nervous sys-
tem, producing either physiological or psychological results. Plant fami-
lies producing alkaloids include the Apocynaceae, Berberidaceae, Fabaceae,
Papaveraceae, Ranunuculaceae, Rubiaceae, and Solanaceae. Some well-known
alkaloids include caffeine, cocaine, ephedrine, morphine, nicotine, and
quinine.
Glycosides
Glycosides are compounds that combine a sugar, usually glucose, with an
active component. While there are many types of glycosides, some of the
most important groups of potentially poisonous glycosides include the
cyanogenic, cardioactive, anthraquinone, coumarin, and saponin glyco-
sides.
Cyanogenic glycosides are found in many members of the Rosaceae and
are found in the seeds, pits, and bark of almonds, apples, apricots, cherries,
peaches, pears, and plums. When cyanogenic glycosides break down, they
release a compound called hydrogen cyanide.
Two other types of glycosides, cardioactive glycosides and saponins,
feature a steroid molecule as part of their chemical structure. Digitalis, a
cardioactive glycoside, in the right amounts can strengthen and slow the
heart rate, helping patients who suffer from congestive heart failure. Other
cardioactive glycosides from plants such as milkweed and oleander are
highly toxic. Saponins can cause severe irritation of the digestive system
and hemolytic anemia. Anthraquinone glycosides exhibit purgative activi-
ties. Plants containing anthraquinone glycosides include rhubarb (Rheum
species) and senna (Cassia senna).
Household Plants
Many common household plants are poisonous to both humans and ani-
mals. One family of popular household plants that can cause problems
is the Araceae, the philodendron family, including plants such as philo-
dendron and dieffenbachia. All members of this family, including these
plants, contain needlelike crystals of calcium oxalate that, when ingested,
cause painful burning and swelling of the lips, tongue, mouth, and throat.
This burning and swelling can last for several days, making talking and
even breathing difficult. Dieffenbachia is often referred to by the common
name of dumb cane, because eating it makes people unable to talk for a few
days.
491
Poisonous plants
Foxgloves, a common ornamental

garden flower, produce cardiac
glycosides with strong physiological
effects on heart muscle. Although
highly toxic, if processed in the right
amounts these glycosides can
strengthen and slow the heart rate,
helping patients who suffer from
congestive heart failure.
(PhotoDisc)
Landscape Plants
Many landscape plants are also poisonous. For example, the yew (genus
Taxus), commonly planted as a landscape plant, is deadly poisonous.
Children who eat the bright red aril, which contains the seed, are poisoned
by the potent alkaloid taxine. Yews are poisonous to livestock as well, caus-
ing death to horses and other cattle. Death results from cardiac or respira-
tory failure.
Other poisonous landscape and garden plants include oleander, rhodo-
dendrons, azaleas, hyacinths, lily of the valley, daffodils, tulips, and Star-
of-Bethlehem. Many legumes are also toxic, including rosary pea, lupines,
and wisteria. Castor bean plant, a member of the family Euphorbiaceae, pro-
duces seeds that are so toxic that one seed will kill a child and three seeds
are fatal to adults. The toxin produced by the seeds is called ricin, which
many scientists consider to be the most potent natural toxin known.
Arrow Poisons
Toxic plant and animal products have been used for thousands of years in
hunting, executions, and warfare. Usually the poisonous extracts were
smeared on arrows or spears. The earliest reliable written evidence for
these uses comes from the Rigveda from ancient India. Arrow poisons come
in many different varieties, and most rain-forest hunters have their own se-
cret blend. South American arrow poisons are generically called curare.
492
Poisonous plants
There are more than seventy different plant species used in making arrow
poisons. Two of the main arrow poison plants are woody vines from the
Amazon: Strychnos toxifera and Chondodendron tomentosum. Some types of
curare have proven medically useful. They are used as muscle relaxants in
surgery, which lessens the amount of general anesthetic needed. A plant
called Strychnos nux-vomica from Asia yields the poison strychnine, a stim-
ulant of the central nervous system.
In ancient times, toxic plant products were also commonly used in exe-
cutions. Many people were expert, professional poisoners in the ancient
world. They could select a poison that would take days or even months to
take effect, thus ensuring, for example, that an unfaithful spouse or lover
would not suspect the reason for his or her lingering illness. On occasions
when a more rapid result was required, a strong dose or more powerful
poison could be prescribed.
Poison Ivy
Toxicodendron radicans, commonly known as poison ivy, is well known for
causing contact dermatitis. Poison ivy is a member of the Anacardiaceae, or
cashew family, and is a widespread weed in the United States and south-
ern Canada. It grows in a variety of habitats: wetlands, disturbed areas,
and the edges of forests. It has many forms, appearing as either a shrub or a
woody vine which will grow up trees, houses, fences, and fence posts. It
has alternate leaves with three leaflets, forming the basis of the old saying
“Leaves of three, let it be.” After poison ivy flowers, it develops clusters of
white or yellowish-white berries. Related species are poison oak, western
poison oak, and poison sumac, which some scientists consider to be differ-
ent types of poison ivy.
Roughly half the world’s population is allergic to poison ivy. Very sensi-
tive people develop a severe skin rash; about 10 percent of the people who
are allergic require medical attention after exposure. The chemical com-
pound causing the allergic reaction is called urushiol, a resin found in all
parts of the plant. Urushiol is so potent that in some individuals, just one
drop produces a reaction. Inhaling smoke from burning poison ivy can re-
sult in eye and lung damage. For some people, mere contact with the
smoke from burning poison ivy can trigger a reaction. Urushiol lasts for-
ever; in herbaria, dried plants one hundred years old have given unlucky
botanists contact dermatitis.
Carol S. Radford

Metabolites; Pheromones; Poisonous animals; Predation.
493
Poisonous plants

Burrows, George E., and Ronald J. Tyrl. Toxic Plants of North America. Ames:
Iowa State University Press, 2001.
Levetin, Estelle, and Karen McMahon. Plants and Society. 2d ed. Boston:
WCB/McGraw-Hill, 1999.
Lewis, Walter H., and Memory P. F. Elvins-Lewis. Medical Botany: Plants Af-
fecting Man’s Health. New York: John Wiley and Sons, 1997.
Simpson, Beryl B., and Molly Conner Ogarzaly. Economic Botany: Plants in
Our World. 3d ed. Boston: McGraw-Hill, 2000.
494
POLLINATION
Types of ecology: Community ecology; Physiological ecology
Pollination—the transfer of pollen from anther to stigma in flowering plants or

from male cone to ovules in gymnosperms—accounts for a wide variety of ecologi-
cal interactions in communities of organisms.
P ollination is the process, in sexually reproducing plants (both angio-

sperms and gymnosperms), whereby the male sperm and female egg
are joined via transfer of pollen (male microspore). If the anthers and stig-
mas of the plants involved have the same genetic makeup or they are pro-
duced on the same plant, the type of pollination is called self-pollination. If
anthers and stigmas are from plants with different genetic makeups, the
type of pollination is called cross-pollination.
Self-pollination is efficient because pollen from the anther of a flower
can be transferred easily onto the stigma of the same flower, owing to the
proximity of the two parts. On the other hand, cross-pollination is risky be-
cause the transfer of pollen involves long distances and precise destina-
tions, both of which depend on animal pollinators. In areas with few ani-
mal pollinators, the opportunities for cross-pollination may be greatly
reduced (one of the many reasons that preserving biological diversity is an
important ecological issue).
In spite of the risk associated with cross-pollination, most flowers have
mechanisms that promote this kind of pollination. Cross-pollination in-
creases the likelihood that offspring are vigorous, healthy, fertile, and able
to survive even if the environment changes. Self-pollination leads to off-
spring that are less vigorous, less productive, and more subject to inbreed-
ing depression (weakening of the offspring as a result of inbreeding).
When certain consumers forage among plants for food, they often come
in contact with flowers. Many insects and other animals become dusted
with pollen, and in the course of their travel they unintentionally but effec-
tively bring about pollination. Throughout the evolutionary history of
flowering plants, many pollinators have coevolved with plants. Coevolu-
tion occurs when the floral parts of a plant and the body parts and behavior
of the pollinators become mutually adapted to each other, thereby increas-
ing the effectiveness of their interaction. In many instances, the relation-
ship between the plant and pollinator has become highly specialized, re-
sulting in mutualism, which is interaction where both organisms benefit
from each other.
495
Pollination
In the case of pollination by animals, the pollinator receives a reward

from the flower in the form of food. When the pollinator moves on, the
plant’s pollen is transferred to another plant. The adaptations between the
flower and its pollinators can be intricate and precise and may even in-
volve force, drugs, deception, or sexual enticement. In flowering plants,
pollination is mostly due to insects or wind, but birds, bats, and rodents
also act as pollinators for a number of plants.
Insects
Insect pollination occurs in the majority of flowering plants. There is no
single set of characteristics for insect-pollinated flowers, because insects
are a large and diverse group of animals. Rather, each plant may have a set
of reproductive features that attracts mostly a specific species of insect. The
principal pollinating insects are bees, although many other kinds of insects
act as pollinators, including wasps, flies, moths, butterflies, ants, and bee-
tles.
Bees have body parts suitable for collecting and carrying nectar and
pollen. Their chief source of nourishment is nectar, but they also collect
pollen for their larvae. The flowers that bees visit are generally brightly
colored and predominantly blue or yellow—rarely pure red, because red
appears black to bees. The flowers they visit often have distinctive mark-
ings that function as guides that lead them to the nectar. Bees can perceive
Honeybees are well known for their

contribution to plant propagation,
carrying pollen from flower to flower.
This relationship has coevolved over
time; the bees are dependent on the
flowers, and vice versa. (PhotoDisc)
496
Pollination
ultraviolet (UV) light (a part of the spectrum not visible to humans), and
some flower markings are visible only in UV light, making patterns per-
ceived by bees sometimes different from those seen by humans. Many bee-
pollinated flowers are delicately sweet and fragrant.
Moth- and butterfly-pollinated flowers are similar to bee-pollinated
flowers in that they frequently have sweet fragrances. Some butterflies can
detect red colors, and so red flowers are sometimes pollinated by them.
Many moths forage only at night; the flowers they visit are usually white
or cream-colored because these colors stand out against dark backgrounds
in starlight or moonlight. With their long mouthparts, moths and butter-
flies are well adapted for securing nectar from flowers with long, tube-
shaped corollas (the petals collectively), such as larkspur, nasturtium, to-
bacco, evening primrose, and amaryllis.
The flowers pollinated by beetles tend to have strong, yeasty, spicy, or
fruity odors. They are typically white or dull in color, in keeping with
the diminished visual sense of their pollinators. Although some beetle-
pollinated flowers do not secrete nectar, they furnish pollen or other foods
which are available on the petals in special storage cells.
Birds
Birds and the flowers that they pollinate are also adapted to each other.
Birds do not have a highly developed sense of smell, but they have a keen
sense of vision. Their flowers are thus frequently bright red or yellow and
usually have little, if any, odor. The flowers are typically large or are part of
a large inflorescence. Birds are highly active pollinators and tend to use up
their energy very rapidly. Therefore, they must feed frequently to sustain
themselves. Many of the flowers they visit produce copious quantities of
nectar, assuring the birds’ continued visitation. The nectar is frequently
produced in long floral tubes, which prevent most insects from gaining ac-
cess to it. Examples of bird-pollinated flowers are red columbine, fuchsia,
scarlet passion flower, eucalyptus, hibiscus, and poinsettia.
Bats and Rodents

Bat-pollinated flowers are found primarily in the tropics, and they open
only at night, when the bats are foraging. These flowers are dull in color,
and like bird-pollinated flowers, they are large enough for the pollinator
to insert part of its head inside. The plants may also consist of ball-like in-
florescences containing large numbers of small flowers whose stamens
readily dust the visitor with pollen. Bat-pollinated flowers include ba-
nanas, mangoes, kapok, and sisal. Like moth-pollinated flowers, flowers
that attract bats and small rodents open at night. Mammal-pollinated flow-
497
Pollination
ers are usually white and strongly scented, often with a fruity odor. Such
flowers are large, to provide the pollinators enough pollen and nectar to
fulfill their energy requirements. The flowers are also sturdy, to bear the
frequent and vigorous visits of these small mammals.
Orchid Pollinators
The orchid family has pollinators among bees, moths and butterflies, and
beetles. Some of the adaptations between orchid flowers and their polli-
nators are extraordinary. Many orchids produce their pollen in little sacs
called pollinia, which typically have sticky pads at the bases. When a bee
visits such a flower, the pollinia are usually deposited on its head. In some
orchids, the pollinia are forcibly “slapped” on the pollinator through a trig-
ger mechanism within the flower. In some orchids, a petal is modified so
that it resembles a female wasp or bee. Male wasps or bees emerge from
their pupal stage before the females and can mistake the orchids for poten-
tial mates. They try to copulate with these flowers, and while they are do-
ing so, pollinia are deposited on their heads. When the wasps or bees visit
other flowers, the pollinia are caught in sticky stigma cavities.
When moths and butterflies pollinate orchids, the pollinia become at-
tached to their long tongues by means of sticky clamps instead of pads.
The pollinia of certain bog orchids become attached to the eyes of the fe-
male mosquitoes that pollinate them. After a few visits, the mosquitoes are
blinded and unable to continue their normal activities (a good example of a
biological control within an ecosystem).
Among the most bizarre of the orchid pollination mechanisms are those
whose effects are to dunk the pollinator in a pool of watery fluid secreted
by the orchid itself and then permit the pollinator to escape underwater
through a trap door. The route of the insect ensures contact between the
pollinia and stigma surfaces. In other orchids with powerful narcotic fra-
grances, pollinia are slowly attached to the drugged pollinator. When the
transfer of pollinia has been completed, the fragrance abruptly fades away,
and the insect recovers and flies away.
Wind and Water

Wind pollination is common in those plants with inconspicuous flowers,
such as grasses, poplars, walnuts, alders, birches, oaks, and ragweeds.
These plants lack odor and nectar and are, hence, unattractive to insects.
Furthermore, the petals are either small or absent, and the sex organs are
often separate on the same plant. In grasses, the stigmas are feathery and
expose a large surface to catch pollen, which is lightweight, dry, and easily
blown by the wind. Because wind-pollinated flowers do not depend on an-
498
Pollination
imals to transport their pollen, they do not invest in the production of re-
wards for their visitors. However, they have to produce enormous quanti-
ties of pollen. Wind pollination is not efficient because most of the pollen
does not end up on the stigmas of appropriate plants but on the ground,
bodies of water, and in people’s noses (a major cause of allergic reactions).
Wind pollination is successful in cases where a large number of individu-
als of the same species grow fairly close together, as in grasslands and co-
niferous forests.
Water pollination is rare, simply because fewer plants have flowers that
are submerged in water. Such plants include the sea grasses, which release
pollen that is carried passively by water currents. In some plants, such as
the sea-nymph, pollen is threadlike, thus increasing its chances of coming
in contact with stigmas. In eelgrass, the entire male flower floats.
Danilo D. Fernando
See also: Adaptations and their mechanisms; Animal-plant interactions;

Coevolution; Communities: ecosystem interactions; Communities: struc-
ture; Reproductive strategies; Symbiosis.

Barth, Friedrich G. Insects and Flowers: The Biology of a Partnership. Prince-
Proctor, Michael, Peter Yeo, and Andrew Lack. The Natural History of Polli-
nation. Portland, Oreg.: Timber Press, 1996.
Raven, Peter H., Ray F. Evert, and Susan E. Eichhorn. Biology of Plants. 6th
ed. New York: W. H. Freeman/Worth, 1999.
499
POLLUTION EFFECTS
Pollutants in soil, water, and atmosphere have created enormous problems for the
living world. Destroyed habitats and polluted food sources and drinking water for
animals have caused deformations in animal growth, development, and reproduc-
tion, as well as a shortening of life span, all of which contribute to an accelerated
decrease in biodiversity and the extinction of more species.
D uring the last decade of the twentieth century, the ecological prob-
lems predicted by environmental scientists decades previously began
to accelerate in a variety of ways. These included the human population
explosion, food imbalances, inflation brought about by energy resource
scarcity, acid rain, toxic and hazardous wastes, water shortages, major soil
erosion, a punctuated ozone layer, and greenhouse effects. As a result of
pollution, decreases in biodiversity and the extinction of both plant and
animal species has accelerated. The burning and cutting of thousands of
square miles of rain forests not only destroyed habitats for numerous ani-
mal species but also caused irreversible damage to ecosystems and cli-
mates. Industrialization and the expansion of the human population had
left relatively few places on earth undisturbed. Heavy dependence upon
fossil fuels for energy and synthetic chemicals has resulted in the dumping
of millions of metric tons of nonnatural compounds and chemicals into the
environment.
Recurrent drought and famine in Africa testify to human mischief to-
ward Mother Nature. The well-being of animals as well as humans will not
be protected against the ecological consequences of human actions by re-
maining ignorant of those actions. Effective measures taken to reduce pol-
lution and protect natural resources and the environment first come with a
recognition of these problems. The ignorance and inaction of ordinary citi-
zens will lead to disastrous consequences for the environment, threatening
humanity’s very existence.
Sources and Types of Pollution

Among the primary sources of pollution are agrichemicals such as fertiliz-
ers, insecticides, fungicides, and herbicides. The application of excess
chemical fertilizers applied to soil hampers natural cycling of nutrients,
depletes the soil’s own fertility, and destroys the habitats of thousands of
small animals residing in the soil. Farm runoff carries priceless topsoil, ex-
500
Pollution effects
pensive fertilizer, and animal manure into rivers and lakes, where these
potential resources become pollutants and cause eutrophication and the
subsequent death of fish and other wildlife.
In the city, water pours from sidewalks, rooftops, and streets, picking
up soot, silt, oil, heavy metals, and garbage. It races down gutters into
storm sewers, carrying household pollutants from cleaning solutions to
prescription medications, and a weakly toxic soup gushes into the nearest
stream, river, or ocean. Many of these chemicals also seep into the ground,
causing contamination of groundwater.
Plants and factories manufacturing these chemical products are another
source of pollutants and contamination. Burning fossil fuels releases
greenhouse gases, carbon dioxide, and methane. Coupled with deforesta-
tion in many regions of the world, carbon dioxide concentration in the at-
mosphere has steadily climbed, from 290 parts per million in 1860 to 370
parts per million in 1990, a more than 25 percent rise due to industrializa-
tion. The resultant global warming will have far-reaching effects on plants,
animals, and humans in ways still not understood. Acid rain, a result of
overcharging the atmosphere with nitric oxides and sulfur dioxide (two
gases also released by burning of fossil fuels), has increased the acidity of
soil and lakes to levels at which many organisms cannot survive. The most
acidic rain is concentrated in the Northeast of the United States. In New
York’s Adirondack Mountains, for instance, acid rain has made about a
third of all the lakes and ponds too acidic to support fish. First, much of the
food web that sustains the fish was destroyed. Clams, snails, crayfish, and
insect larvae die first, then amphibians, and finally fish. The detrimental
effect is not limited to aquatic animals. The loss of insects and their larvae
and small aquatic animals has contributed to a dramatic decline in the pop-
ulation of black ducks that feed on them. The result is a crystal-clear lake,
beautiful but dead.
Another serious problem created by the chemical industry is ozone de-
pletion. Chlorofluorocarbon (CFC) compounds contain chlorine, fluorine,
and carbon. Since their development in the 1930’s, these compounds were
widely used as coolants in refrigerators and air conditioners, as aerosol
spray propellants, as agents for producing Styrofoam, and as cleansers for
electronic parts. These chemicals are very stable and for decades were con-
sidered to be safe. Their stability, however, turned out to be a real problem.
They were in gaseous form and rose into the atmosphere. There, the high
energy level of ultraviolet (UV) light breaks them down, releasing chlorine
atoms, which in turn catalyzes the breakdown of ozone to oxygen gas. As a
result of the decline of ozone and the punctuation of the ozone layer, UV
radiation has risen by an average of 8 percent per decade since the 1970’s.
501
Pollution effects
Air Pollutant Emissions by Pollutant and Source, 1998
Volatile
Sulfur Nitrogen Organic Carbon Lead
1
Source Particulates Dioxide Oxides Compounds Monoxide (tons)
Fuel Combustion
(stationary sources)
Electric utilities 302 13,217 6,103 54 417 68
Industrial 245 2,895 2,969 161 1,114 19
Other fuel combustion 544 609 1,117 678 3,843 416
Residential 432 127 742 654 3,699 6
Subtotal 1,091 16,721 10,189 893 5,374 503
Industrial processes
Chemical and allied 65 299 152 396 1,129 175
product manufacturing
Metals processing 171 444 88 75 1,495 2,098
Petroleum and related 32 345 138 496 368 NA
industries
Other 339 370 408 450 632 54
Subtotal 607 1,458 786 1,417 3,624 2,327
Solvent utilization 6 1 2 5,278 2 NA
Storage and transport 94 3 7 1,324 80 NA
Waste disposal and 310 42 97 433 1,154 620
recycling
Highway vehicles
Light-duty gas vehicles 56 130 2,849 2,832 27,039 12
and motorcycles
Light-duty trucks 40 99 1,917 2,015 18,726 7
Heavy-duty gas vehicles 8 11 323 257 3,067 —
Diesels 152 86 2,676 222 1,554 NA
Subtotal 257 326 7,765 5,325 50,386 19
Off highway2 461 1,084 5,280 2,461 19,914 503
Miscellaneous3 31,916 12 328 786 8,920 NA
Total emissions 34,742 19,647 24,454 17,917 89,454 3,972
Source: Adapted from U.S. Environmental Protection Agency, National Air Pollutant
Emission Trends, 1900-1998, EPA-454/R-00-002. From Statistical Abstract of the United
States: 2001 (Washington, D.C.: U.S. Bureau of the Census, 2001).
Note: In thousands of tons, except as indicated.
— Represents or round to zero.
NA Not available
1
Represents both particulates of less than 10 microns and particulate dust from sources
such as agricultural tilling, construction, mining, and quarrying, paved roads, unpaved
roads, and wind erosion.
2
Includes emissions from farm tractors and other farm machinery, construction equipment,
industrial machinery, recreational marine vessels, and small general utility engines such
as lawn mowers.
3
Includes emissions such as from forest fires and other kinds of burning, various
agricultural activities, fugitive dust from paved and unpaved roads, and other
construction and mining activities, and natural sources.
502
Pollution effects
This depletion of the ozone layer poses a threat to humans, animals, plants,
and even microorganisms.
Long-Range Impacts of Pollution

The degradation of air, land, and water as a result of the release of chemical
and biological wastes has wide-ranging effects on animals. On a large
scale, pollution destroys habitats and produces population crashes and
even the extinction of species. Hazardous chemicals introduced into the
environment sometimes render an environment unfit for life (as at Love
Canal, New York, or Times Beach, Missouri). At the individual level, pollu-
tion causes abnormalities in growth, development, and reproduction. Haz-
ardous chemicals, introduced either intentionally (such as fertilizers, her-
bicides, and pesticides) or through neglect (as with industrial wastes),
have a variety of detrimental, sometimes devastating effects on animals.
They affect the metabolism, growth and development, reproduction, and
average life spans of many species.
A few examples will illustrate the effects of chemical pollution on ani-
mals. In the 1940’s, the new insecticide dichloro-diphenyl-trichloroethane
(DDT) was regarded as a miracle. It saved millions of lives in the tropics by
killing the mosquitoes that spread deadly malaria. DDT saved millions
more lives with increased crop yields resulting from DDT’s destruction of
insect pests. This miraculous pesticide, however, turned out to be a long-
lasting nemesis to many species of wildlife and the environment. In the
United States, ecologists and wildlife biologists during the 1950’s and
1960’s witnessed a stunning decline in the populations of several preda-
tory birds, especially fish-eaters, such as bald eagles, cormorants, ospreys,
and brown pelicans. The population decline drove the brown pelican and
bald eagle close to extinction. In 1973, the U.S. Congress passed the Endan-
gered Species Act, which banned the use of DDT. The once-threatened spe-
cies have somewhat recovered since. In the mid 1950’s, the World Health
Organization used DDT on the island of Borneo to control malaria. DDT
entered food webs through a caterpillar. Wasps that fed on caterpillars
were first destroyed. Gecko lizards that ate the poisoned insects accumu-
lated high levels of DDT in their bodies. Both geckos and the village cats
that ate the geckos died of DDT poisoning. The rat population exploded
with its natural enemy, cats, eliminated. The village was then threatened
with an outbreak of plague, carried by the uncontrolled rats.
Although DDT has been banned in much of the world, there is a grow-
ing concern over the effects of a number of chlorinated compounds. These
chemicals, described as “environmental estrogens,” interfere with normal
sex hormone functions by mimicking the effects of the hormone estrogen
503
Pollution effects
or enhancing estrogen’s potency. High levels of chlorinated compounds,

such as dioxin and polychlorinated biphenyls (PCBs), in the Great Lakes
have led to a sharp decline in populations of river otters and a variety of
fish-eating birds, including the newly returned bald eagles. These chemi-
cals are also the cause of deformed offspring, or eggs that never hatch. In
Florida’s Lake Apopka, a spill of chlorinated chemicals in 1980 led to a 90
percent drop in the birthrate of the lake’s alligators. These are only a few
examples of the detrimental effects on various animals by synthetic chemi-
cals.
Air Pollution
Air pollution leads to acid rain and the greenhouse effect, as well as damage
to the ozone layer. Acid rain drops out of the skies onto areas at great dis-
tances from the source of the acids and destroys forests and lakes in sensi-
tive regions. As a result, fish populations are dwindling or being elimi-
nated in lakes and streams by a lower pH caused by acid deposition. The
strongest evidence comes from data collected from the past twenty-five to
forty-five years in Adirondack lakes and in Nova Scotia rivers. Studies
during this period clearly show declines in acid-sensitive species. Similar
results were obtained from analyzing fish population and water acidity in
Maine, Massachusetts, Pennsylvania, and Vermont. The consensus is that
fish populations would be eliminated if the surface waters acidify to be-
tween pH 5.0 to pH 5.5. The effects of acid rain on other animals are indi-
rect, either through the dwindling fish population (as a food source for
other animals) or stunted forest growth (disturbance to habitats).
The effect of global warming on the animal kingdom is also a serious
and complex issue. As global temperature rises, ice caps in polar regions
and glaciers melt, ocean waters expand in response to atmospheric warm-
ing, and thus the sea level elevates. The expected sea level rise will flood
coastal cities and coastal wetlands. These threatened ecosystems are hab-
itats and breeding grounds for numerous species of birds, fish, shrimp,
and crabs, whose populations could be severely diminished. The Florida
Everglades will virtually disappear if the sea level rises two feet. The im-
pact of global warming on forests could be profound. The distribution of
tree species is exquisitely sensitive to average annual temperature, and
small changes could dramatically alter the extent and species composition
of forests. This in turn could dramatically alter the population distribution
of animals, and hence biodiversity.
The effect of the punctuated ozone layer on animals is yet to be fully un-
derstood. It is known that the high energy level of UV radiation can dam-
age biological molecules, including the genetic material deoxyribonu-
504
Pollution effects
cleic acid (DNA), causing mutation. In small quantities, UV light helps the
skin of humans and many animals produce vitamin D and causes tan-
ning. However, in large doses, UV causes sunburn and premature aging of
skin, skin cancer, and cataracts, a condition in which the lens of the eye be-
comes cloudy. Due to UV radiation’s ability to penetrate, even animals
covered by hair and thick fur cannot escape from these detrimental effects.
Ozone damage costs U.S. farmers over $2 billion annually in reduced crop
yields. All who depend on forestry and agriculture may bear a much
higher cost if the emission of pollutants that destroy ozone are not regu-
lated soon.
Possible Remedies
The various types of pollution all have serious effects on the plant and ani-
mal species that share this planet. It is all too easy to document the impacts
of pollution on human health and ignore their effects on the rest of the liv-
ing world. Any possible remedies to alleviate these problems should start
with education, the realization of these problems at an individual as well
as a global level. The tasks seem to be insurmountable, and no organiza-
tion, no country can do it alone. It takes willingness to accept short-term in-
convenience or economic sacrifice for long-term benefit. A couple of exam-
ples serve to illustrate what can be done to alleviate the problems of
pollution.
Synthetic chemical pollutants that are poisoning both people and wild-
life could be largely eliminated without disrupting the economy, as re-
ported in a study published in 2000 by the Worldwatch Institute, a Wash-
ington, D.C.-based environmental organization. The report presents strong
evidence from three sectors that are major sources of these pollutants—
paper manufacturing, pesticides, and PVC plastics—to show that nontoxic
options are available at competitive prices. Agricultural pollution can be
mitigated, significantly reduced, or virtually eliminated through the use of
proper regulation and economic incentives. Farmers from Indonesia to
Kenya are learning how to use less of various chemicals while boosting
yields. Since 1998, all farmers in China’s Yunnan Province have elimi-
nated their use of fungicides, while doubling rice yields, by planting more
diverse varieties of the grain. In most, if not all, cases, the question is
not whether it is possible to alleviate the pollution of the environment;
rather, it is whether we realize the urgency and/or are willing to take a
high road to do it. For the common well-being of generations to come,
better approaches have to be taken to preserve the environment and biodi-
versity.
Ming Y. Zheng
505
Pollution effects
See also: Acid deposition; Biological invasions; Biomagnification; Biopes-

ticides; Deforestation; Eutrophication; Genetically modified foods; Inte-
grated pest management; Invasive plants; Ocean pollution and oil spills;
Ozone depletion and ozone holes; Pesticides; Phytoplankton; Slash-and-
burn agriculture; Waste management.

Brown, Lester. State of the World 2000. New York: W. W. Norton, 2000.
Hill, Julia B. The Legacy of Luna: The Story of a Tree, a Woman, and the Struggle
to Save the Redwoods. San Francisco: HarperSanFrancisco, 2000.
Johnson, Arthur H. “Acid Deposition: Trends, Relationships, and Effects.”
Environment 28, no. 4 (May, 1986): 6-11, 34-39.
Lippmann, Morton, ed. Environmental Toxicants: Human Exposures and Their
Health Effects. 2d ed. New York: John Wiley & Sons, 2000.
Sampat, Payal. Deep Trouble: The Hidden Threat of Groundwater Pollution.
Washington, D.C.: Worldwatch Institute, 2000.
506
POPULATION ANALYSIS
Many animal populations are becoming threatened or endangered, primarily due

to loss of suitable habitat. Population analysis enables biologists to examine the
factors which lead to declines in animal populations and thus is important in the
management of wild species.
A population is a group of organisms belonging to the same species

that occur together in the same time and place. Population analysis is
the study of biological populations, with the specific intent of understand-
ing which factors are most important in determining population size. Pop-
ulations can change over time. They increase or decrease in size, and their
change in size can depend on a wide variety of factors. For example, a
wildlife biologist might be interested in studying the population of porcu-
pines that inhabits a hemlock forest or the population of bark beetles that
lives on a particular tree.
Population analysis is the study of biological populations, with the spe-
cific intent of understanding which factors are most important in deter-
mining population size. Factors such as the per capita rates of birth and
death, the population density, age structure, and sex ratio all contribute to
determine population size. Understanding how these factors interact to in-
fluence population size is critical if biologists hope to manage populations
of organisms at sustainable levels for hunting or fishing and if conserva-
tion biologists hope to prevent populations from going extinct.
Discrete vs. Continuous Populations

In order to conduct a population analysis, one must first determine whether
the population of interest is best understood as discrete or continuous. A
discrete population is one in which important events such as birth and
death happen during specific intervals of time. A continuous population is
one in which births, deaths, and other events take place continuously
through time. Many discrete populations are those with nonoverlapping
generations. For example, in many insect populations, the adults mate and
lay eggs, after which the adults die. When the juveniles achieve adulthood,
their parental generation is no longer living. In contrast, most continuous
populations also have overlapping generations. For instance, in antelope
jackrabbits (Lepus alleni), females may give birth at any time during the
year, and members of several generations occur together in space and time.
507
Population analysis
Mathematical Models
The dynamics of animal populations are affected by a wide variety of de-
mographic factors, including the population birthrate, death rate, sex ra-
tio, age structure, and rates of immigration and emigration. In order to un-
derstand the effects of these factors on a population, biologists use popula-
tion models. A model is an abstract representation of a concrete idea. The
representation created by the model boils the concrete idea down into a
few critical components. By building and examining population models,
population analysts investigate the relative importance of different factors
to the dynamics of a given population.
A basic mathematical model of population size is as follows:
Nt+1 − Nt + B − D + I − E (equation 1)
where Nt+1 equals the population size after one time interval, Nt equals the
total number of individuals in the population at the initial time, B equals
the number of births, D equals the number of deaths, I equals the number
of immigrants into the population, and E equals the number of emigrants
leaving the population. This simple model boils population size down to
just four factors, B, D, I, and E. This model is not meant to be a true or pre-
cise representation of the population; rather, it is meant to clarify the im-
portance of the factors of birth, death, immigration, and emigration on
population size. To use the same model to examine the rate of growth of a
population through time, it can be rearranged as follows:
Nt+1 − Nt = B − D + I −E (equation 2)
That is, the increase or decrease in the population size between time inter-
vals t and t + 1 is reflected by the number of births, deaths, immigrants, and
emigrants.
When population biologists choose to focus specifically on the impor-
tance of birth and death in population dynamics, population models are
simplified by temporarily ignoring the effects of immigration and emigra-
tion. In this case, the degree of change in the population between time in-
tervals t and t + 1 becomes:
Nt+1 − Nt = B − D (equation 3)
It is usually safe to assume that the total number of births (B) and deaths
(D) in a population is a function of the total number of individuals in the
population at the time, Nt. For example, if there are only ten females in a
508
Population analysis
population at time t, it would be impossible to have more than ten births in

the population. More births and deaths are possible in larger populations.
If B equals the total number of births in the population, then B is equal to the
rate at which each individual in the population gives birth, times the total
number of individuals in the population. Likewise, the total number of
deaths, D, will be equal to the rate at which each individual in the popula-
tion might die times the total number of individuals in the population. In
other words:
B = bNt and D = dNt (equation 4)
where b and d represent the per capita rate of birth and death, respectively.
Given this understanding of B and D, the original model becomes
Nt+1 − Nt = (bNt) − (dNt)

or
Nt+1 − Nt = (b − d)Nt (equation 5)
It would be useful to find a variable that can represent per capita births and
deaths at the same time. Biologists define r as the per capita rate of increase
in a population, which is equal to the difference between per capita births
and per capita deaths:
r=b−d (equation 6)
Thus, the equation that examines the changes in population size between
time intervals t and t + 1 becomes:
Nt+1 − Nt = rNt (equation 7)
A numerical example works as follows. In a population that originally

has 1,000 individuals, a per capita birthrate of 0.1 birth per year and a per
capita death rate of 0.04 death per year, the net change in the population
size between the year t and t + 1 would be:
r = 0.1 − 0.04 = 0.06
Nt+1 − Nt = 0.06(1000) = 60
In other words, the population would increase by sixty individuals over

the course of one year.
509
Population analysis
Continuous Populations
This model works for populations in which events take place during dis-
crete units of time, such as a population of squirrels in which reproduction
takes place at only two specific times in a single year. In contrast, many
populations are continuously reproductive. That is, at any given time, any
female in the population is capable of reproducing. When these conditions
are met, time is viewed as more fluid than discrete, and the population ex-
hibits continuous growth. Models of population growth are slightly differ-
ent when births and deaths are continuous rather than discrete. One way
to imagine the difference between a population with continuous rather
than discrete growth is to imagine a population in which each time interval
is infinitesimally small. When these conditions are met, the model for pop-
ulation growth becomes:
δN/δt = rN (equation 8)
where δN/δt represents the changes in numbers in the population over

very short time intervals. The per capita rate of increase (r) can now also be
called the instantaneous rate of increase because the population is one with
minute time intervals.
Choosing the Right Model

How does a population biologist select the best model? Which model is
best depends on exactly what it is that a scientist is trying to understand
about a population. In the first model presented above (equation 1), the dif-
ferent effects of birth, death, immigration and emigration can be compared
relative to one another. In the second model, the effects of immigration
and emigration are ignored and the effects of birth and death are summa-
rized into one constant called the per capita rate of increase (equations 7
and 8). If the scientist is trying to understand the cumulative effects of B, D,
I, and E on the population, then equation 1 would represent a good model.
On the other hand, if the scientist is trying to understand how births and
deaths influence the net changes in population size, equation 7 or 8 would
be a better model.
When dealing with a continuous rather than a discrete population,
equation 8 represents the rate of population growth as a function of per ca-
pita births and per capita deaths in the population. Equation 8 represents a
population that is growing exponentially without bound. In other words,
regardless of the population size at any given time, the per capita rate of in-
crease remains the same. It would be reasonable to assume that per capita
rates of increase can actually change with changes in overall population
510
Population analysis
size. For example, in a population of bark beetles inhabiting the trunk of a

tree, many more resources are available to individual beetles when the
population is small. Resources must be shared between more and more in-
dividuals as the population size increases, which can result in changes to
the per capita rate of increase. A model of population growth that incorpo-
rates the effect of overall population density on the per capita rate of in-
crease might look like this:
δN/δt = r(1 − N/K)N (equation 9)
where K is equal to the carrying capacity, the maximum number of individ-

uals in the population that there are adequate resources to support. The
per capita rate of increase in equation 9 is not simply r by itself, but be-
comes r(1 − N/K). The per capita rate of increase is a function of rates of
birth and death scaled by the population size and the carrying capacity of
the habitat. If the population is very large relative to the number of individ-
uals that the habitat can support, then N ≈ K, and the expression (1 − N/K)
becomes approximately equal to 0. When so, equation 9 takes the follow-
ing form:
δN/δt = r(0)N = 0 (equation 10)
and the rate of population growth is zero. In other words, the population
has ceased growing. On the other hand, if the population is very small rela-
tive to the number of individuals the habitat can support, then N << K and
the expression (1 − N/K) becomes approximately equal to 1. When so,
equation 9 takes the form
δN/δt = r(1)N = rN (equation 11)
and the rate of population growth remains a function of the rates of birth
and death, but not the population size or carrying capacity. Thus, equation
9 represents what is called density-dependent growth.
Sex Ratio and Age Structure

The model set forth in equation 9 takes into account only those ways in
which births, deaths, and population density relative to carrying capacity
influence population growth. Sometimes it is helpful to understand how
other factors such as the sex ratio and age structure in a population influ-
ence rates of growth. For example, deer hunters are not always allowed to
take equal numbers of bucks and does from a population. Similarly, fisher-
511
Population analysis
men are often restricted in the size of fish they are allowed to keep when
fishing. These wildlife management and population analysis restrictions
on the sex and size of animals that can be hunted arise from the fact that
both age and sex can influence population growth rates. Models that incor-
porate the effects of age structure and population sex ratios will not be cov-
ered here. Suffice it to say that a population that consists mostly of young
individuals yet to reproduce will grow more quickly than an equally sized
population of mostly older individuals who have finished reproducing.
Similarly, a population with a highly skewed sex ratio that has many more
males than females will not grow as quickly as a population of equal size in
which the number of males and females is equal.
Erika L. Barthelmess
See also: Biodiversity; Biogeography; Clines, hybrid zones, and intro-

gression; Demographics; Extinctions and evolutionary explosions; Gene
flow; Genetic diversity; Genetic drift; Human population growth; Nonran-
dom mating, genetic drift, and mutation; Population fluctuations; Popula-
tion genetics; Population growth; Reproductive strategies; Speciation; Spe-
cies loss.

Gardali, Thomas, et al. “Demography of a Declining Population of War-
bling Vireos in Coastal California.” The Condor 102, no. 3 (August, 2000):
601-609.
Hastings, Alan. Population Biology: Concepts and Models. New York:
Hedrick, Philip W. Population Biology: The Evolution and Ecology of Popula-
tions. Boston: Jones and Bartlett, 1984.
Johnson, Douglas H. “Population Analysis.” Research and Management
Techniques for Wildlife and Habitats, edited by Theodore A. Bookhout. 5th
ed. Bethesda, Md.: The Wildlife Society, 1994.
512
POPULATION FLUCTUATIONS
The simplest realistic models of population growth produce populations that rise to
some level and then stay there. These models cannot produce the complicated array
of fluctuations observed in natural populations. Fluctuations vary in period from a
few weeks to many decades and can reach sufficient amplitude to threaten popula-
tions (and entire species) with extinction.
T he number of organisms making up a population is never constant; it

always changes over time. The populations of some species change in
predictable or cyclical ways, whereas populations of other species fre-
quently exhibit seemingly unpredictable and noncyclic changes. Fluctua-
tions in population size may be caused by changes in the population’s en-
vironment; for example, seasonal changes in temperature or moisture
produce seasonal fluctuations in population size. Resource limitations
may produce density-dependent reductions in the growth rate of a popu-
lation, which, if the reduction is not instantaneous, can result in oscilla-
tions in population size. Interactions with other species also produce pop-
ulation fluctuations; mathematical models of predator-prey systems
typically produce oscillations in the abundance of both predators and prey.
Finally, natural or anthropogenic disturbances often reduce the size of a
population, which then either recovers its former abundance over time or
declines further to local (or global) extinction.
The Time Scales of Population Fluctuations

Population fluctuations occur over many different time scales. On a geo-
logic time scale (occurring over millions of years), species arise, increase to
some level of abundance, and finally become extinct. These long-term pat-
terns of species abundance provide a background for understanding pop-
ulation fluctuations that occur over ecological time (over days, weeks,
years, or centuries). Fluctuations on these briefer time scales draw most of
the attention of ecologists interested in population dynamics.
Many species of animals, including numerous insects and several small
vertebrates, exhibit a more or less annual life cycle, characterized by in-
creasing numbers and higher levels of activity during the summer (or wet
season) and by dormancy or decreasing numbers during the winter (or dry
season). Even highly mobile animals, such as birds, exhibit a strong sea-
sonal pattern of abundance, if viewed from a local perspective; in North
513
Population fluctuations
America, for example, most songbirds migrate to more tropical latitudes in

the fall and to temperate latitudes in the spring, thereby producing a yearly
cycle of abundance in each location. Yearly cycles of abundance are pre-
dictable and easily explainable in terms of seasonal patterns of tempera-
ture, moisture, and sunlight. Of more interest to ecologists are population
fluctuations that appear to be random or unpredictable from year to year
or those fluctuations that occur out of synchrony with climatic cycles.
Regular Fluctuations
Nonseasonal fluctuations are of two main types: those that exhibit more or
less regular cycles of abundance over several years and those that seem to
fluctuate irregularly or noncyclically. A three- to four-year cycle of abun-
dance is characteristic of several species of mice, voles, and other rodents
found in far northern latitudes. Probably the best-known example of this
type of cycle is that observed in lemming species in the northern tundra of
Europe and North America. Lemming populations exhibit very high den-
sities every three to four years, with such low densities in the intervening
years that they are difficult to locate and study. This boom-or-bust cycle is
apparently caused by alternating selection regimes. When lemmings are
rare, high reproductive capacity and nonaggressive social behavior are fa-
vored, and the population grows rapidly. As the growing population be-
comes more crowded, aggressive individuals are favored, because they
can hold territories, secure mates, and protect offspring better than passive
individuals. The aggressive interactions, however, inhibit reproductive ca-
pacity, increase mortality attributable to fighting and infanticide, and ex-
pose more lemmings to predation as subordinate individuals are forced by
dominants to occupy more marginal habitats. The behavioral changes that
occur in response to crowding apparently persist for some time even as the
density declines, so that aggressive interactions and a depressed birthrate
continue until the lemming population reaches very low levels. Finally,
passive individuals with high reproductive rates are again favored, and
the cycle repeats.
Although the breeding cycles of many predators, including snowy
owls, weasels, and foxes, are tied to lemming abundance, it appears that
the regular fluctuation of lemming populations is a product of crowding
and resource limitation rather than of a classical predator-prey cycle; that
is, there is no tight coupling between the population fluctuations of lem-
mings and those of their predators. There is, however, a tight coupling
between the population cycles of the snowshoe hare and the Canadian
lynx. Beginning in 1800, the Hudson’s Bay Company kept records of furs
produced each year. Both the hare and the lynx showed a regular ten-year
514
cycle, with the peaks in lynx abundance occurring about a year behind
the hare’s peak abundances. Since the hare is a major food source for the
lynx in northern Canada, it is logical to assume that this is a coupled oscil-
lation of population sizes, precisely as predicted by classical predator-prey
theory.
Some regular cycles of abundance appear to have evolved as a means
of avoiding predation rather than being a direct reduction caused by pre-
dation. There is a periodicity in the populations of cicadas and locusts.
The hypothesized explanation is that predators cannot reproduce rapidly
enough to increase their population sizes quickly in response to the sud-
den availability of a large food supply. When millions of adult cicadas
appear above ground for a few weeks after surviving for seventeen years
as nymphs in the soil, predators cannot possibly consume them all: No
predator could specialize on adult cicadas unless it also had a seventeen-
year cycle.
Several northern bird populations (such as crossbills, grosbeaks, and
waxwings) fluctuate dramatically, in some years rising to several times
their usual levels. This fluctuation may be a response to changing habitat
quality. These bird populations always produce as many eggs as food
availability and their natural fecundity allow, even though many offspring
will not survive. In a good year, a higher proportion of the offspring sur-
vives, and the population experiences an irruption, often leading to intense
competition and consequent expansion of the range of the population. In
subsequent years, population size returns to preirruption levels. Thus,
these fluctuations are entirely consistent with normal density-dependent
processes responding to a fluctuating environment.
Irregular Fluctuations
Population fluctuations that occur irregularly or noncyclically often ap-
pear to be responses to natural disturbances rather than to density-depen-
dent processes or predator-prey relationships. For example, blue grouse
persists at a relatively low level of abundance in coniferous forests until a
fire occurs. The species rapidly increases in number following a fire and
gradually diminishes again as the forest regenerates over the next several
decades.
The population fluctuations of some species are not easily attributed to
disturbance or to any other single cause. For example, swarming locusts
typically remain at low abundance in a restricted area for several years;
then, apparently without warning, they may increase more than a hun-
dredfold and swarm over large areas, consuming large amounts of veg-
etation. The locust outbreak lasts for several years, then the population
515
declines as rapidly as it initially increased. In the early part of the twenti-

eth century, it was discovered that locusts exhibit two phases: a solitary
phase, corresponding to low abundance, and a gregarious phase, corre-
sponding to an outbreak. While it is still not known how locusts trans-
form from one phase to another, it is clear that several stages are involved
and that weather conditions seem to initiate a transformation. Moisture
seems to be the most important determinant, because of its influence
on nymph development and survival, on egg development, and on preda-
tor abundance, but wind and plant nitrogen levels have also been impli-
cated. Furthermore, it appears that environmental conditions are only ef-
fective in inducing a phase transformation if a certain concentration of
locusts already exists and if the existing locusts are adequately sensitive to
crowding.
Measuring Fluctuations
There are two parts to the study of population fluctuation: detecting and
measuring the pattern of the fluctuation and identifying the underlying
causes of the fluctuation. In general, any method designed for measuring
population size can be used repeatedly over time to detect fluctuations
in the population. Reference to a specialized textbook on ecological sam-
pling techniques is strongly recommended when using any of these meth-
ods, in order to assure validity of the sampling for subsequent statistical
analysis. The mark-recapture method is commonly used with animal pop-
ulations. There are many variants of this technique, but they all involve
capturing and marking some number of individuals, then releasing them;
after some time period appropriate to the study, a second sample is cap-
tured and the proportion of marked individuals in the second sample
(those that are “recaptured”) is recorded. This proportion is used to esti-
mate the size of the population at the time when the individuals were origi-
nally marked.
The quadrat method is used primarily with plants and other sessile or-
ganisms. Plots (called quadrats) are laid out, either randomly or in some
pattern; all individuals within the plots constitute a sample. Quadrats are
usually square, but any regular shape may be used. The appropriate size of
each quadrat depends on the sizes of organisms to be sampled and on their
spatial distribution. If nondestructive sampling techniques are used, the
same quadrats may be sampled repeatedly; otherwise, new quadrats must
be established for each sampling episode.
A variety of plotless techniques are available for sessile organisms, in
lieu of the quadrat method. These techniques were developed to eliminate
some of the uncertainties associated with selecting proper quadrat size
516
and location. Most plotless methods locate points on the ground, then
measure distances to nearby organisms; each plotless technique identifies
the individuals to be measured in a slightly different way.
None of these techniques is adequate by itself to identify the origin or
cause of any fluctuation in population size. Experimental manipulation of
a population is necessary to elucidate the underlying mechanisms and de-
termining factors. Populations of small, rapidly reproducing species (such
as species of Paramecium or Daphnia) can be manipulated in the laboratory,
and hypothesized causes of fluctuation can be tested under controlled con-
ditions. This has been done primarily to develop theoretical predictions re-
garding environmental conditions (such as temperature, moisture, and hu-
midity), resource limitations and fluctuations, and the effects of predators
and competitors.
Identifying Causes of Fluctuation

The most interesting examples of population fluctuation, however, occur
over spatial and temporal scales too large to handle in the laboratory. Their
underlying mechanisms must be elucidated in the field. Because suites of
factors typically produce the complex patterns of population fluctuation
observed in nature, an effective field study must include all relevant fac-
tors. Generally, the most effective studies have been those that have sought
to understand the complete life history of a species. Superb examples in-
clude the long-term studies of the wolves of Isle Royal National Park, by
David Mech and colleagues, and the equally ambitious studies of the griz-
zly bears of Yellowstone National Park and surrounding areas by Frank
and John Craighead and their many coworkers.
Most equilibrium models of population dynamics are capable of pro-
ducing regular oscillations that mimic the patterns observed in nature. If
the model parameters are properly manipulated, many of these models
can produce apparently random fluctuation. More sophisticated models
have been constructed that incorporate a mathematical equivalent of ran-
dom environmental fluctuation, although they usually still assume that a
population has a tendency to stabilize and that environmental change sim-
ply prevents stabilization. The underlying assumption of almost all these
models is that species normally exist at equilibrium. This assumption is
consistent with the long-held belief that there is a “balance of nature”—
that species exist in harmony with their environment.
If an entire species is considered, perhaps the assumption of equilib-
rium is warranted, at least for extended periods; yet at the level of the pop-
ulation, fluctuation is the rule—indeed, it may be that extreme fluctuation
is the rule. As noted earlier, many populations fluctuate so markedly that
517
they often disappear; they are reestablished only by colonization from

large populations within dispersal range. If populations become too small
or too isolated from one another, this colonization cannot occur. Addi-
tionally, because small populations are more subject to extinction associ-
ated with fluctuation, there is additional risk of species extinction if only
small populations remain.
The problem of extinction is severe, since habitat destruction is occur-
ring at an unprecedented rate on a global scale. The fragments of intact
habitat that remain because of inaccessibility or preservation efforts con-
tain populations that are smaller and more isolated than in the past. If an
isolated population fluctuates markedly, resulting in its extinction from a
habitat fragment, its replacement by recolonization is unlikely. Further-
more, the genetic variation maintained by a complex population structure
within a species is reduced. As the genetic variation within a species is lost,
the ability of a species to respond to environmental change is reduced, and
extinction of the species is more likely.
Ultimately, if populations normally fluctuate severely enough that they
can be expected to become extinct at frequent intervals, then effective con-
servation requires the maintenance of pathways for exchange and dis-
persal of individuals among populations within a species. It also requires
the preservation of the largest population size possible to allow for normal
fluctuation without extinction.
Alan D. Copsey

dom mating, genetic drift, and mutation; Population analysis; Population
genetics; Population growth; Reproductive strategies; Speciation; Species
loss.

Begon, Michael, Martin Mortimer, and David J. Thompson. Population
Ecology: A Unified Study of Animals and Plants. 3d ed. Cambridge, Mass.:
Blackwell Science, 1996.
Associates, 2001.
dance. 4th ed. New York: HarperCollins College Publishers, 1994.
Krebs, Charles J., and J. H. Myers. “Population Cycles in Small Mammals.”
Advances in Ecological Research 8, 1974): 267-399.
518
Mech, L. David. The Wolf: The Ecology and Behavior of an Endangered Species.
Garden City, N.Y.: Doubleday, 1970.
Smith, Robert Leo. Elements of Ecology. 4th ed. San Francisco: Benjamin/
Cummings, 2000.
Wohrmann, K., and S. K. Jain, eds. Population Biology: Ecological and Evolu-
tionary Viewpoints. New York: Springer-Verlag, 1990.
519
POPULATION GENETICS
Population genetics is the analysis of genes and genetic traits in populations to de-
termine how much variability exists, what maintains the variability, how selection
(natural or controlled) affects a population, and what the mechanisms of evolution
are.
C lassical genetics deals with the rules of genetic transmission from par-
ents to offspring, developmental genetics deals with the role of genes
in development, and molecular genetics looks at the molecular basis of ge-
netic phenomena. Population genetics uses information from all three
fields and helps explain why populations are so variable, why some harm-
ful traits are common, why most animals and plants reproduce sexually,
how evolution works, why some animals are altruistic in a cutthroat
world, and how new species arise.
Mutations and Natural Selection

Simple observation reveals that animals are highly variable. Some dogs are
big, others small; some wiry, others big boned; some long-haired, others
curly; some with special talents such as herding or retrieving, others with
none; and some with diseases or defects, others normal. All of these are the
result of various genes combined with environmental influences. Unless
an animal is an identical twin, no one else shares that individual’s geno-
type and no one ever will. Population genetics looks at variability in a pop-
ulation and examines its sources and the forces that maintain it.
Variability can come from genetic mutations. For example, about one
child in ten thousand is born with dominant achondroplasia (short-legged
dwarfism). Some children with the trait inherit the condition from an af-
fected parent, but most have normal parents. They are therefore the result
of a new mutation. Many mutations are deleterious and are eventually
eliminated from the population by the lowered survival or fertility rates of
those who have the mutation, but while they remain in the population,
they add to its variability. Occasionally, a seemingly harmful mutation per-
sists, for example, the gene that causes sickle-cell disease, a severe disease
characterized by red blood cells that become sickle shaped in certain labo-
ratory tests. The causative gene is recessive, meaning that two copies are
needed to produce the anemia, but the disease is very common in some
parts of Africa. The harmful, anemia-causing gene persists in the popula-
520
Population genetics
tion because if a person has only one gene with the trait rather than two,
that gene confers resistance to malaria, the major cause of debility in that
part of the world.
Although the genes in these two examples have large and conspicuous
effects, the great majority of mutations and the great bulk of genetic vari-
ability in the population are the result of a large number of genes with indi-
vidually small effects, often detected only through statistics. The variabil-
ity of quantitative traits such as size is due mainly to the cumulative action
of many individual genes, each of which produces its small effect. The av-
erage size stays roughly constant from generation to generation because
individuals who are too large or too small are at a disadvantage. However,
such individuals continuously arise from new mutations.
The driving force in evolution is natural selection, that is, the differen-
tial survival and fertility of different genotypes. New mutations occur con-
tinuously. Most of these are harmful, although usually only mildly so, but
a small minority are beneficial. The rules of Mendelian inheritance ensure
that the genes are thoroughly scrambled every generation. Natural selec-
tion acts like a sieve, retaining those genes that produce favorable pheno-
types in the various combinations and rejecting others. Such a process, act-
ing over eons of time, has produced the variety and specific adaptations
that can be found throughout the animal kingdom.
The Hardy-Weinberg Law and Selfish Genes

Although evolutionary progress is the result of natural selection, most se-
lection does not accomplish any systematic change. Most selection is di-
rected at maintaining the status quo—eliminating harmful mutations,
keeping up with transitory changes in the environment, and eliminating
statistical outliers (extremes of variation). Most of the time, evolutionary
change is very slow.
In most populations, mating is essentially random in that mates do not
choose each other because of the genes they carry. There are exceptions, of
course, but for the most part, random mating can be assumed. This permits
a great simplification known as the Hardy-Weinberg rule. This rule says
that if the proportion of a certain gene, say A, in the population is p and of
another, say a, is q, then the three genotypes AA, Aa, and aa are in the pro-
2 2
portions p , 2pq, and q , respectively. (Remember that p and q are fractions
between zero and one.) This is a simple application of elementary proba-
bility and the binomial theorem. Furthermore, after a few generations of
random mating, genotypes at different loci also equilibrate, which means
that the frequency of a composite genotype is the product of the frequen-
cies at the constituent loci. The reason that this is so useful is that the num-
521
Population genetics
ber of genotypes is enormous, but a population can be characterized by a

much smaller number of gene frequencies.
Genotypes are transient, but genes may persist unchanged for many
generations. This has led the great theorists of population genetics, J. B. S.
Haldane and R. A. Fisher in England and Sewall Wright in the United
States, to make the primary units the frequency of individual genes and
develop theories around this concept, making free use of the simple conse-
quences of random mating. Such a gene-centered view has been described
by scholar Richard Dawkins as the “selfish gene.” A population can be
thought of as a collection of genes, each of which is maximizing its chance
of being passed on to future generations. This causes the population to be-
come better adapted because those genes that improve adaptation have
the best chance of being perpetuated.
Kin Selection and Selfish Genes

An extension of this notion is kin selection. The concept holds that, to the
extent that behavior is determined by genes, individuals should be protec-
tive of close relatives because relatives share genes. The fact that brothers
and sisters share half their genes should lead a brother to be half as con-
cerned with his sister’s survival and reproduction as with his own. Many
evolutionists believe that altruistic behavior in various animals, including
humans, is the result of kin selection. The degree of self-sacrifice to protect
a close relative is proportional to the fraction of shared genes. Parents regu-
larly make sacrifices for their children, and this is what evolutionary the-
ory would predict.
One way in which populations depart from random mating is inbreed-
ing, the mating of individuals more closely related than if they were ran-
domly chosen. Related individuals share one or more ancestors, hence an
inbred individual may get two copies of an ancestral gene, one through
each parent. In this way, inbreeding increases the proportion of homozy-
gotes. Because many deleterious recessive genes are hidden in the popula-
tion, inbreeding can have a harmful effect by making genes homozygous.
Similarly, if the population is subdivided into local units, mating mostly
within themselves, these local units will be more homozygous than if the
entire population mated at random. Small subpopulations will be more
subject to purely random fluctuations in gene frequencies known as ran-
dom genetic drifts. Therefore, subdivisions of a population often differ sig-
nificantly, particularly with respect to unimportant genes.
Views of Evolution: Gene-Centered vs. Genetic Drift

The gene-centered view of evolution is not always accepted. Some evolu-
522
Population genetics
tionists believe that it is simplistic to view an individual as a bag of genes,

each trying to perpetuate itself. They emphasize that genes often interact in
complicated ways, and that a theory that deals with only average gene ef-
fects is incomplete. Modern theories of evolution take such complications
into account.
This different viewpoint has led to a major controversy in evolution,
one that has not yet been settled. Wright emphasized that many well-
adapted phenotypes depend on genes that interact in very specific ways;
two or more genes may be individually harmful but when combined
produce a beneficial effect. He argued that selecting genes on the basis
of average effects cannot produce such combined effects. He believed
that a population subdivided into many partially isolated units provides
an opportunity for such interactions. An individual subpopulation, by
random drift, might chance upon such a happy gene combination, in
which case the whole population can be upgraded by migrants from this
subpopulation. Whether evolutionary advance results from gene interac-
tions in subpopulations, from mass selection in largely unstructured popu-
lations, or from a combination of both is a question that remains unre-
solved.
Population genetics theory, along with the techniques of molecular ge-
netics, has greatly deepened our knowledge of historical evolution. Most
students of evolution are familiar with tree diagrams of common ancestry
that show, for example, birds and mammals branching off from early an-
cestors. In the past these had to be constructed using external phenotypes
and fossils. These techniques for measuring the relatedness of different
species and determining their ancestral relations have been replaced by
DNA sequencing, which produces much surer results. It has long been sus-
pected that genes can persist for very long evolutionary periods, modify-
ing slightly to perform new, often related but sometimes quite different
functions. This belief has been confirmed repeatedly by molecular analy-
sis. The similarity of the DNA sequences between some plant and animal
genes is so great as to leave no doubt that they were both derived from a
common ancestral gene a billion or more years ago.
Neutral Mutation and Sexual Reproduction

Most gene mutations have very small effects, and the smaller the effect, the
less likely it is to be noticed. Molecular techniques have enabled scientists
to detect changes in DNA without regard to the traits they cause or
whether they have any effect at all. The Japanese geneticist Motoo Kimura
has advanced the idea that most evolution at the DNA level is not the result
of natural selection but simply the result of mutation and chance, a concept
523
Population genetics
termed neutral mutation. In vertebrates, especially in mammals, most of

the DNA has no known function. The functional genes make up a very
small fraction of the total DNA. Many scientists believe that most DNA
evolution outside the genes—and some within—is the result of changes
that are so nearly neutral as to be determined by chance. How large a role
random drift plays in the evolution of changes in functional proteins is still
not certain.
A few animals and a large number of plants reproduce asexually. In-
stead of reproducing by using eggs and sperm, the progeny are car-
bon copies of the parent. Asexual reproduction has obvious advantages.
If females could reproduce without males, producing only female off-
spring like themselves, reproduction would be twice as efficient. How-
ever, despite its inherent inefficiency, sexual reproduction is the rule, un-
doubtedly because of the gene-scrambling process that sex produces. The
ability of a species to produce and try out countless gene combinations
confers an evolutionary advantage that outweighs the cost of males.
Another advantage of gene scrambling is that it permits harmful muta-
tions to be eliminated from the population in groups rather than individu-
ally.
Population genetics is also concerned with the processes by which new
species arise. Scientists believe that a population somehow becomes di-
vided into two or more isolated groups, separated perhaps by a river,
mountain range, or other geographical barrier. Each group then follows its
own separate evolutionary course, and the groups’ dissimilar environ-
ments accentuate their differences. Eventually the two groups evolve so
many differences that they are no longer compatible. The products of
interspecies crosses, or hybrids, often do not develop normally or are ster-
ile (like the mule). Sometimes the two species do not mate because they are
so different.
Research Methods
Population genetics involves theory, observation, and experiment. Popula-
tion genetics examines how genes are influenced by mutation, selection,
population size, migration, and chance. Scientists develop mathematical
models that embody these theories and compare the results obtained using
the models with data from laboratory experiments or field observations.
These genetic models have become more and more sophisticated to take
into account complex gene interactions and increasingly realistic popula-
tion structures. The models are further complicated by efforts to account
for random processes. Often the mathematical geneticist relies on comput-
ers to perform complex analyses and computations.
524
Population genetics
One of the simpler models, which makes the assumption that mating is
random, is the Hardy-Weinberg principle. If the proportion of gene A in
the population is p and that of gene a is q, then the three genotypes AA, Aa,
and aa are in the proportions p2, 2pq, and q2, respectively. The proportion of
Aa is 2pq rather than simply pq because this genotype represents two com-
binations, maternal A with paternal a and paternal A with maternal a. This
principle can be used to predict the frequency of persons with malaria re-
sistance from the incidence of sickle-cell anemia. If one-tenth of the genes
are sickle-cell genes and the other nine-tenths are normal, the frequency of
two genes coming together to produce an anemic child is 0.1 × 0.1, or 0.01.
The frequency of those resistant to malaria, who have one normal and one
sickle-cell gene, is 2 × 0.1 × 0.9, or 0.18. A slight extension of the calculation
(using the rates of malaria infection and death from the disease) can be
used to estimate the death rate from malaria. Another mathematical model
can be formed based on the molecular genetics theory of neutral mutation.
A neutral mutation, because it is not influenced by natural selection, has an
expected rate of evolution that is equal to the mutation rate. Mathematical
models embodying this theory are used to quantitatively predict what will
happen in an experiment or what an observational study will find and act
as a test of the theory. Neutral mutation theory is quite complicated and re-
quires advanced mathematics.
Observational population genetics consists of studying animals and
plants in nature. Evolution rates are inferred from the fossil record.
Field observations can determine the frequency of genes in different
geographical areas or environments. The frequency of self- and cross-
pollination can often be observed directly. Increasingly, DNA analysis,
which can detect relationships or alterations that are not visible, is be-
ing used to support field observations. For example, molecular mark-
ers have been used to determine parentage and relationship. DNA analy-
sis revealed that certain birds that do not reproduce but care for the
progeny of others are in fact close relatives, consistent with kin selection
theory.
Increasingly, population genetics has begun to rely on experimentation.
Plants and animals can be used to study the process of selection, but to
save time and reduce costs, most laboratory experiments involve small,
rapidly reproducing organisms such as the fruit fly, Drosophila. Some of the
most sensitive selection experiments have involved the use of a chemostat,
a container in which a steady inflow of nutrients and steady outflow of
wastes and excess population permits a population to maintain a stable
number of rapidly growing organisms, usually bacteria. These permit very
sensitive measurements of the effects of mutation. Evolutionary studies
525
Population genetics
that would require eons if studied in large animals or even mice can be
completed in a very short time.
Uses of Population Genetics

The greatest intellectual value of population genetics has been to provide a
theory of evolution that is explanatory, quantitative, and predictive. Popu-
lation genetics places knowledge of mutation, gene action, selection, in-
breeding, and population structure in a unified framework. It brings to-
gether Charles Darwin’s theory of evolution by natural selection, Gregor
Mendel’s laws of inheritance, and molecular genetics to create a coherent
picture of how evolution took place and is still occurring.
Population genetics has provided explanations for variability in a pop-
ulation, the prevalence of sexual rather than asexual reproduction, the
origin of new species, and behavioral traits such as altruism. It has also
provided an understanding of why some harmful diseases are found in
the population. Population genetics has been used in animal and plant
breeding to create rational selection programs. Using quantitative models,
the results of various selection schemes can be compared and the best one
chosen.
A particularly telling example of a situation in which population genet-
ics predicted an outcome that has become painfully obvious is the devel-
opment of resistance to insecticides, herbicides, and antibiotics. As people
used these products more and more, the insects, weeds, and bacteria they
were trying to eliminate developed resistance, and new products had to
be developed to replace those rendered ineffective. The development of
resistance represents evolution by natural selection that took place not
over hundreds or thousands of years but in just a few years. Probably
the most problematic area of resistance is antibiotics because some treat-
able diseases are again threatening to become beyond the ability of medi-
cine to cure. A major challenge to ecologists, microbiologists, physicians,
and population geneticists is how to deal with the increasingly difficult
problem of disease-producing microorganisms that are resistant to antibi-
otics.
James F. Crow

dom mating, genetic drift, and mutation; Population analysis; Population
fluctuations; Population growth; Reproductive strategies; Speciation; Spe-
cies loss.
526
Population genetics

Crow, James F. Basic Concepts in Population, Quantitative, and Evolutionary
Genetics. New York: W. H. Freeman, 1986.
Dawkins, Richard. The Blind Watchmaker. New York: W. W. Norton, 1986.
_______. The Selfish Gene. Rev. ed. Oxford, England: Oxford University
Press, 1999.
Falconer, Douglas S. Introduction to Quantitative Genetics. 4th ed. New York:
John Wiley & Sons, 1996.
Fisher, R. A. The Genetical Theory of Natural Selection. Rev. ed. New York: Do-
ver, 1958.
Haldane, J. B. S. The Causes of Evolution. Reprint. Ithaca, N.Y.: Cornell Uni-
Hartl, Daniel. A Primer of Population Genetics. 3d ed. Sunderland, Mass.:
Hartl, Daniel, and Andrew Clark. Principles of Population Genetics. 3d ed.
Kimura, Motoo. The Neutral Theory of Molecular Evolution. Cambridge, En-
gland: Cambridge University Press, 1985.
Maynard Smith, John. The Theory of Evolution. Cambridge, England: Cam-
Wright, Sewall. Evolution and the Genetics of Populations. 4 vols. Chicago:
University of Chicago Press, 1968-1978.
527
POPULATION GROWTH
Populations typically grow when they are found on sites with abundant resources,
and biologists have developed two models to describe growth. In exponential
growth, the population is exposed to ideal conditions, and new individuals are
added at an ever-increasing rate. Logistic growth recognizes that resources are
eventually depleted, however, and that the population density ultimately stabilizes
at some level, which is defined as the carrying capacity.
I n nature, organisms of a particular species rarely occur by themselves.

Instead, they usually exist with other individuals of the same species. Bi-
ologists use the term “population” to refer to an aggregation of organisms
of a given species that live in the same general location at the same time. In
some cases, populations can be well defined, such as herds of cattle or
flocks of geese. In other cases, the population is not well defined, often be-
cause several species may be found in the same location. For example, a
meadow may contain intermingled populations of several species, includ-
ing daisies, timothy grass, earthworms, and grasshoppers.
Biologists have studied populations for many years. Many of those
studies have been conducted to answer three separate but interrelated
questions. First, how many individuals are there in a given population at a
particular time? Second, how do those numbers change from one time to
another? Third, what environmental factors are responsible for any popu-
lation increases or decreases? Studies have shown that for most species of
plants, animals, and microbes, the number of individuals in the population
changes over time. Some populations increase steadily, other populations
decrease, and still others fluctuate. Thus, populations, when viewed over
time, are generally dynamic rather than static.
Population Behaviors
Most populations change so much through time because there is constantly
turnover among individuals. That is, new individuals are constantly being
born, hatched, or germinated, while others die. Moreover, animals are also
able to enter a population by immigration and leave by emigration. Since
the number of births and new immigrants hardly ever exactly matches the
number of deaths and emigrants, the dynamic nature of populations
should not be a surprise. Because changes in population size are common
in nature, biologists have tried to understand the changes that are ob-
528
Population growth
served. One approach has been to model populations; the model is a sim-
plified graphical or mathematical summary of the actual changes that are
occurring in the species of interest. The relationship between a model and
the actual population that it represents is similar to that between a map
and the area of land that it represents. Because modeling is such an impor-
tant aspect of population biology, biologists who study population must
often have a good background in mathematics.
Perhaps the simplest mode of population behavior is the difference
equation, which states that the number of individuals in a population at
some specified time in the future is equal to the number at present, plus the
number of births, minus the number of deaths, plus the number of new im-
migrants, minus the number of emigrants. Thus, by knowing how many
individuals are on a site at a given time and knowing the usual number of
births, deaths, immigrants, and emigrants, one can predict the number of
individuals on the site at some future time.
Obviously, the number of births, deaths, immigrants, and emigrants
will vary from one place to another and from one time to another. For ex-
ample, on a site with abundant food and space and with favorable physical
conditions for growth and development, births and immigration will be
much greater than deaths and emigration. Thus, the population will in-
crease. Conversely, if food or space is limiting or if the physical conditions
are more severe, losses to the population through death and emigration
will equal or exceed gains through birth and immigration. Thus, the popu-
lation will remain constant or decline.
Biologists often are concerned about what happens in extreme con-
ditions, because such conditions define the limits within which the pop-
ulation normally operates. When conditions are very bad, a population
normally declines rapidly, often to the point of local extinction; when con-
ditions are very good, a population will increase. That increase is attribut-
able to the fact that each individual normally has the capacity to produce
many offspring during its lifetime. For example, a woman could produce
more than forty children if she conceived every time that she was fertile.
Other individual organisms, particularly many invertebrates and plants,
can produce hundreds of thousands of offspring in their lifetime.
Birth and Death Rates

At least three different traits influence the reproductive output of a given
species. The first is the number of offspring per reproductive period (ele-
phants produce only one child at a time, whereas flies can lay thousands of
eggs). The second is the age at first reproduction (most dogs can reproduce
when less than three years old, whereas humans do not usually become
529
Population growth
fertile until they reach an age of thirteen or fourteen). The third is the num-
ber of times that an individual reproduces in its lifetime (salmon spawn,
that is, lay eggs, only once before they die, whereas chickens lay eggs re-
peatedly). Even under ideal conditions, death must also be considered
when examining population growth. Nearly all organisms have a maxi-
mum life span that is determined by their innate physiology and cannot be
exceeded, even if they are supplied with abundant food and kept free from
disease.
Population biologists frequently express birth and death in the form of
rates. This can be done by counting the number of new births and deaths in
a population during a predetermined period of time and then dividing by
the number of individuals in the population. That will give the per capita
(per individual) birth and death rates. For example, suppose that during
the course of a year there were thirty births and fifteen deaths in a popula-
tion of one thousand individuals. That per capita birthrate would be 0.030
and the per capita death rate would be 0.015.
Next, one can subtract the death rate from the birthrate to find the per
capita rate of population growth. That rate should be greatest under ideal
conditions, when the birthrate is greatest and the death rate is least. That
per capita rate of population growth is called the “maximal intrinsic rate of
increase” or the “biotic potential” by population biologists, and it is a very
important attribute. It is often symbolized as rmax or referred to as “little r.”
Normally, rmax is considered an inherent feature of a species. As one might
expect, it varies greatly among different types of organisms. For example,
rmax, expressed per year, is 0.02-1.5 for birds and large mammals, 4-50 for in-
sects and small invertebrates, and as high as 20,000 for bacteria.
Exponential Growth
By knowing the intrinsic rate of increase and the number of organisms in a
population, one can predict much about the behavior of a population un-
der ideal conditions. The rate at which the population grows is merely the
intrinsic rate of increase (rmax) multiplied by the number of individuals in
the population. For example, suppose that there are ten individuals in a
population whose annual rmax is 2. That population would increase by an
annual rate of twenty (which would be a healthy increase). Next, suppose
that one returned to that population at some later time when the popula-
tion was fifty individuals. At that point, the annual rate of population in-
crease would be one hundred new individuals (which would be an even
healthier increase). If the rate of increase were measured when the popula-
tion reached five hundred, the annual rate of increase would be one thou-
sand individuals.
530
Population growth
Under such circumstances, the population would keep on growing at

an ever-increasing rate. That type of growth is called “exponential growth”
by population biologists, and it typifies the behavior of many populations
when placed under ideal conditions. If the number of individuals in a pop-
ulation undergoing exponential growth is plotted as a function of time, the
curve would resemble the letter J. That is, it would be somewhat flat ini-
tially, but it would curve upward, and at some point it would be almost
vertical. Exponential population growth has been observed to a limited ex-
tent in many different kinds of organisms, both in the laboratory and un-
der field conditions. Examples include protozoans, small insects, and
birds. It should be obvious, however, that no species could behave in this
manner for long. If it did, it would overrun the earth (and indeed the uni-
verse) in a matter of time. Instead, population growth is slowed by limited
resources, accumulated wastes, behavioral stresses, and/or periodic catas-
trophes caused by the environment.
Logistic Growth
Biologists have created a second model to account for the behavior of pop-
ulations under finite resources and have called it logistic growth. If the
number of individuals in a population undergoing logistic growth is plot-
ted as a function of time, the curve would resemble a flattened S shape. In
other words, the curve would initially be flat, but would then curve up-
ward at a progressively faster rate, much like exponential growth. At some
point (called the inflection point), however, the curve would begin to turn
to the right and flatten out. Ultimately, the curve would become horizontal,
indicating a constant population over time.
An important aspect of pure logistic growth is that the population ap-
proaches, but does not exceed, a certain level. That level is called the carry-
ing capacity, and is represented by the symbol K in most mathematical
treatments of logistic growth. The carrying capacity is the maximum num-
ber of individuals that the environment can support, based on the space,
food, and other resources available. When the number of individuals is
much fewer than the carrying capacity, the population grows rapidly,
much as in exponential growth. As the number increases, however, the rate
of population growth becomes much less than the exponential rate. When
the number approaches the carrying capacity, new population growth vir-
tually ceases. If the population were to increase above the carrying capac-
ity for some reason, there would be a net loss of organisms from the popu-
lation.
There are few studies that have documented logistic growth in nature. It
would be necessary to watch a species in a habitat from the time of its first
531
Population growth
introduction until its population stabilized. Such studies are necessarily of

a very long duration and thus are not normally conducted. Logistic growth
has been found in a number of experimental studies, however, particularly
on small organisms, including protozoans, fruit flies, and beetles.
An important aspect of logistic growth is that, as the population in-
creases, the birthrate decreases and the mortality rate increases. Such ef-
fects may be attributable to reduced space within which the organism can
operate, to less food and other resources, to physiological and behavioral
stress caused by crowding, and to increased incidence of disease. Those
factors are commonly designated as being density-dependent. They are
considered much different from the density-independent factors that typi-
cally arise from environmental catastrophes such as flooding, drought,
fire, or extreme temperatures. For many years, biologists argued about the
relative importance of density-dependent versus density-independent fac-
tors in controlling population size. It is now recognized that some species
are controlled by density-independent factors, whereas others are con-
trolled by density-dependent factors.
Classically, when a species undergoes logistic growth, the population
is ultimately supposed to stabilize at the carrying capacity. Most studies
that track populations over the course of time, however, find that numbers
actually fluctuate. How can such variability be reconciled with the logis-
tic model? On the one hand, the fluctuations may be caused by density-
independent factors, and the logistic equation therefore does not apply. On
the other hand, the population may be under density-dependent control,
and the logistic model can still hold despite the fluctuations. One explana-
tion for the fluctuations could be that the carrying capacity itself changes
over time. For example, a sudden increase in the amount of food available
would increase the carrying capacity and allow the population to grow. A
second explanation relates to the presence of time lags. That is, a popula-
tion might not respond immediately to a given resource level. For example,
two animals in a rapidly expanding population might mate when the
number of individuals is less than the carrying capacity. The progeny, how-
ever, might be born several weeks or months later, into a situation in which
the population has exceeded the carrying capacity. Thus, there would have
to be a decline, leading to the fluctuation.
Approaches to Studying Population Growth

Two main approaches can be used to investigate logistic and exponential
population growth among organisms. One approach involves following
natural populations in the field; the other involves setting up experimental
populations. Each approach has its benefits and drawbacks, and, ideally,
532
Population growth
both should be employed. To study population growth in the field, it is im-

portant to study a species from the time that it first arrives on a site until its
population stabilizes. Thus, any species already present are automatically
eliminated from consideration unless they are brought to local extinction
and a new population is then allowed to recolonize. Population growth
studies can be profitably done on sites that are very disturbed and are be-
ginning to fill up with organisms. Examples would be an abandoned farm
field or strip mine, a newly created volcanic island, or a new body of water.
Moreover, studies could also be done on a species that is purposely intro-
duced to a new site.
In either case, one needs to survey the population periodically to assess
the number of individuals that it contains. The size of the population can
be determined directly or by employing sampling techniques such as
mark-recapture methodology. The number of individuals can then be plot-
ted on a graph (on the y-axis) as a function of time (on the x-axis).
To study population growth in experimental conditions, one sets up an
artificial habitat according to the needs of the species in question. For ex-
ample, investigators have examined population growth in protozoans
(unicellular animals) by growing populations in test tubes filled with food
dissolved in a known volume of water. Others have grown fruit flies in
stoppered flasks. Still others have grown beetles in containers filled with
oatmeal or other crushed grain. In those cases, it was typically necessary to
replenish the food to keep the population going. Whenever the population
was placed into an artificial habitat with a nonrenewable food source, it
would generally consume all the food and then die out.
More detailed experiments can be performed to test whether density-
dependent mortality is occurring. Such experiments would involve setting
up a series of containers with different densities of organisms and then fol-
lowing the mortality of those organisms. In theory, mortality rates should
be highest in containers that have the greatest densities of organisms and
lowest in containers with the sparsest populations. One could also exam-
ine the birthrate in those containers, with the expectation that birthrates
should be highest in the sparsest containers and lowest in those that have
the most organisms. The investigation should be long enough to allow the
population to reach equilibrium at the carrying capacity. For short-lived
organisms such as protozoans or insects, that could take days, weeks, or
months. For longer-lived organisms such as fish or small mammals, one to
several years may be required. For long-lived animals, a truly adequate
study may take decades.
Another consideration in studying exponential and logistic population
growth is that immigration and emigration should be kept to a minimum.
533
Population growth
Thus, organisms that are highly active, such as birds, large mammals, and
most flying insects would be extremely difficult to study. Finally, one can
set up numerous populations and expose each to a slightly different set of
conditions. That would enable the researcher to ascertain which environ-
mental factors are most important in determining the carrying capacity.
For example, populations of aquatic invertebrates could be monitored un-
der a range of temperature, salinity, pH, and nutrient conditions.
Research Applications
Since exponential growth is unrealistic in practical terms for almost all
populations, its scientific usefulness is limited. The concepts derived from
logistic growth, however, have important implications to biologists and
nonbiologists alike. One important aspect of logistic growth is that the
maximum rate of population growth occurs when the population is about
half of the environment’s carrying capacity. When populations are very
sparse, there are simply too few individuals to produce many progeny.
When populations are dense, near the carrying capacity, there is not
enough room or other resources to allow for rapid population growth.
Based on that relationship, those who must harvest organisms can do so at
a rate that allows the population to reestablish quickly.
Those who can apply this concept in their everyday work include wild-
life managers, ranchers, and fishermen. Indeed, quotas for hunting and
fishing are often set in a way that allows for the population to be thinned
sufficiently without depleting it too severely. Unfortunately, there are two
problems that biologists must confront when they try to use the logistic
model to help manage populations. The first is that it is often difficult to es-
tablish the carrying capacity for a given species on a particular site. One
reason is that the populations of many species are profoundly affected by
density-independent factors, as well as by other species, in highly complex
and variable ways. Further, for reasons unclear to most biologists, some
species have a maximum rate of population growth at levels well above or
well below the level (one half of the carrying capacity) that is normally as-
sumed. Thus, the logistic model typically gives only a very rough approxi-
mation for the ideal size of a population. However, the logistic model is
useful because it emphasizes that all species have natural limits to the sizes
of their populations.
Kenneth M. Klemow

534
Population growth
dom mating, genetic drift, and mutation; Population analysis; Popula-

tion fluctuations; Population genetics; Reproductive strategies; Speciation;
Species loss.

Brewer, Richard. The Science of Ecology. 2d ed. Philadelphia: Saunders Col-
lege Publishing, 1994.
Elseth, Gerald D., and Kandy D. Baumgardner. Population Biology. New
York: Van Nostrand, 1981.
Associates, 2001.
Hutchinson, G. Evelyn. An Introduction to Population Ecology. New Haven,
Kormondy, Edward J. Concepts of Ecology. 4th ed. Englewood Cliffs, N.J.:
dance. 5th ed. San Francisco: Benjamin/Cummings, 2001.
Raven, Peter, H., and George B. Johnson. Biology. 5th ed. St. Louis: Times
Mirror/Mosby, 1999.
Wilson, Edward O., and William Bossert. A Primer of Population Biology.
535
PREDATION
The relationships among predators and their prey in natural communities are var-
ied and complex. These interactions provide clues as to how natural populations
regulate one another, as well as to how to preserve and manage exploited popula-
tions more successfully.
P redation is an interaction between two organisms in which one of

them, the predator, derives nutrition by killing and eating the other,
the prey. Obvious examples include lions feeding on zebras and hawks eat-
ing rodents, but predation is not limited to interactions among animals.
Birds that feed on seeds are legitimate predators since they are killing indi-
vidual organisms (embryonic plants) to derive energy. There are a number
of species of carnivorous plants, such as sundews and pitcher plants, that
capture and consume small animals to obtain nitrogen in habitats that oth-
erwise lack sufficient quantities of that nutrient. Most animals that feed on
plants (herbivores) do not kill the entire plant and therefore are not really
predators. Exceptions to this generalization are some insects that reach in-
festation levels, such as gypsy moths or locusts, and can kill the plants
upon which they feed. The majority of herbivore-plant associations, how-
ever, are more properly described as parasite-host interactions in which
the host plant may suffer damage but does not die.
There are special cases in which parasitism and predation may be com-
bined. One of these is the interaction between parasitoid wasps and their
hosts, usually flies. Adult female parasitoids attack and inject eggs into fly
pupae (the resting stage, during which fly larvae metamorphose into
adults), and the larvae of the wasp consume the fly. The adult parasitoid is
therefore a parasite, while the larval wasp acts as a predator.
Predator-Prey Interactions
Predator-prey interactions can be divided into two considerations: the ef-
fects of prey on predators, and the effects of predators on their prey. Preda-
tors respond to changes in prey density (the number of prey in the habitat)
in two principal ways. The first is called numerical response, which means
that predators change their numbers in response to changes in prey den-
sity. This may be accomplished by increasing or decreasing reproduction
or by immigrating to or emigrating from a habitat. If prey density in-
creases, predators may immigrate from other habitats to take advantage of
536
Predation
this increased resource, or those predators already present may produce

more offspring. When prey density decreases, the opposite will occur.
Some predators, which are known as fugitive species, are specialized at
finding habitats with abundant prey, migrating to them, and reproducing
rapidly once they are established.
Cape May warblers are good at finding high densities of spruce bud-
worms (a serious pest of conifers) and then converting the energy from
their prey into offspring. This strategy allows the birds to persist only be-
cause the budworms are never completely wiped out; they are better at
dispersing to new habitats than are the birds.
The second response of predators to changing prey density is called
functional response. The rate at which predators capture and consume
prey depends upon the rate at which they encounter prey, which is a func-
tion of prey density. If the predator has a choice of several prey species, it
may learn to prefer one of them. If that prey is sufficiently abundant, this
situation results in a phenomenon known as switching, the concentration
by the predator on the preferred prey. It may entail a change in searching
behavior on the part of the predator, such that former prey items will no
longer be encountered as frequently.
Animals have evolved a number of defense mechanisms that reduce
their probability of being eaten by predators. Spines on horned lizards,
A hungry bear nabs a salmon for lunch as the fish hurls itself upstream to spawn.
(Corbis)
537
Predation
threatening displays by harmless snakes, camouflage of many cryptic ani-

mals, toxic or distasteful chemicals in insects and amphibians, and simply
rapid movement—all are adaptations that may have evolved in response
to natural selection by predators. A predator that can learn to prefer one
prey item over another is smart enough to learn to avoid less desirable
prey. That capability is the basis for a phenomenon known as aposematism
among potential prey species which are toxic and/or distasteful to their
predators. Aposematic organisms advertise their toxicity by bright color-
ation, making it easy for predators to learn to avoid them, which in
turn saves the prey population from frequent taste-testing. Many species
of insects are aposematic. Monarch butterflies are bright orange with black
stripes, an easy signal to recognize. They owe their toxicity to the milk-
weed plant, which they eat as caterpillars. The plant contains cardiac
glycosides, which are very toxic. The monarch caterpillar is immune to
the poison and stores it in its body so that the adult has a high concentra-
tion of it in its wings. If a bird grabs the butterfly in flight, it is likely to get a
piece of wing first, and this will teach it not to try orange butterflies in the
future.
Some potential prey species that are not themselves toxic have evolved
to resemble those that are; these are called Batesian mimics. Viceroy butter-
flies, which are not toxic, mimic monarchs very closely, so that birds cannot
tell them apart. One limit to Batesian mimicry is that mimics can never get
very numerous, or their predators will not get a strong enough message to
leave them alone. Another kind of mimicry involves mimics that are as tox-
ic as their models. The advantage with this type, Müllerian mimicry, is that
the predator has to learn only one coloration signal, which reduces risk for
both prey populations. In this relationship, the mimic population does not
have to remain at low levels relative to the model population. A third type
of mimicry is more insidious—aggressive mimicry, in which a predator re-
sembles a prey or the resource of that prey in order to lure it close enough
to capture. There are tropical praying mantises that closely resemble or-
chid flowers, thus attracting the bees upon which they prey. There are
some species of fireflies that eat other species of fireflies, using the flashing
light signal of their prey to lure them within range.
The Choice of Prey

What determines predator preference for prey? Since prey are a source of
energy for the predator, it might be expected that predators would simply
attack the largest prey they could handle. To an extent, this choice holds
true for many predators, but there is a cost to be considered. The cost in-
volves the energy a predator must expend to search for, capture, handle,
538
Predation
and consume prey. In order to be profitable, a prey item must yield much
more energy than it costs. Natural selection should favor reduction in ener-
getic cost relative to energetic gain, the basis for optimal foraging theory.
According to this theory, many predators have evolved hunting strategies
to optimize the time and energy spent in searching for and capturing prey.
Some predators, such as web-building spiders and boa constrictors, am-
bush their prey. The low energetic cost of sit-and-wait is an advantage in
environments that provide plentiful prey. If encounters with prey become
less predictably reliable, however, an ambush predator may experience
starvation. Spiders can lower their metabolic energy requirements when
prey is unavailable, whereas more mobile predators, such as boa constric-
tors, can simply shift to active searching. Probably because of the likeli-
hood of facing starvation for extended periods of time, ambush predation
is more common among animals that do not expend metabolic energy to
regulate their body temperatures (ectotherms) than among those that do
(endotherms). Some predators, such as wolves and lions, hunt in groups.
This allows them to tackle larger (more profitable) prey than if they hunted
alone. Solitary hunters generally have to hunt smaller prey.
Natural communities consist of food webs, constructed of links (feed-
ing relationships) among trophic levels. Each prey species is linked to one
or more predators. Most predators in nature are generalists with respect to
their prey. Spiders, snakes, hawks, lions, and wolves all feed on a variety of
prey. Some of these prey are herbivores, but some are themselves preda-
tors. Praying mantises eat grasshoppers (herbivores), but they also eat spi-
ders (carnivores) and each other. Thus, generalist predators have a
bitrophic niche, in that they occupy two trophic levels at the same time.
Predation and Population Fluctuation

It is an open question whether predators and prey commonly regulate each
other’s numbers in nature. There are many examples of cyclic changes in
abundance over time, in which an increase in prey density is followed by
an increase in the numbers of predators, and then the availability of prey
decreases, also followed by a decrease in predators. Are predators causing
their prey to fluctuate, or are prey responding to some other environmen-
tal factor, such as food supply? In the second case, prey may be regulated
by food, and in turn, may be regulating predators, but not the reverse.
Predators can sometimes determine the number of prey species that can
coexist in a habitat. If a predator feeds on a prey species that could
outcompete (competitively exclude) other prey species in a habitat, it may
free more resources for those other species. This relationship is known as
the keystone effect. Empirical studies have indicated that the number of
539
Predation
Although rare, plant predators

exist. This “carnivorous” pitcher
plant is formed with a hollow,
tubular structure that contains
fluid to trap small animals and
insects to obtain nitrogen in
habitats that otherwise lack
sufficient quantities of that
nutrient. (Digital Stock)
prey species in some communities is directly related to the intensity of pre-

dation (numerical and functional responses of predators) such that at low
intensity, few species coexist because of competitive exclusion; at interme-
diate intensity, the diversity of the prey community is greatest; and at high
intensity, diversity decreases because overgrazing begins to eliminate spe-
cies. This intermediate predation hypothesis depends upon competition
among prey species, which is not always the case.
Studying Predation
The central question in the study of predation is: To what extend do preda-
tors and their prey regulate one another? Most studies suggest that preda-
tors are usually food limited, but the extent to which they regulate their
prey is uncertain. It is one thing to observe predators in nature and another
to assess their importance to the dynamics of natural communities. Like
other aspects of ecology, studies of predation can be descriptive, experi-
mental, and/or mathematical.
At the descriptive level, characteristics of both predator and prey popu-
lations are assessed: rates of birth and mortality, age structure, environ-
mental requirements, and behavioral traits. Qualitative and quantitative
information of this type is necessary before predictions can be made about
540
Predation
the interactions between predator and prey populations. General lack of

such information in natural ecosystems is largely responsible for failures at
biological control of pests and management of exploited populations.
Experimental studies of predation involve manipulation of predator
and/or prey populations. A powerful method of testing the importance of
predation is to exclude a predator from portions of its accustomed habitat,
leaving other portions intact as experimental controls. In one such experi-
ment, excluding starfish from marine intertidal communities of sessile in-
vertebrates resulted in domination by mussels and exclusion of barnacles
and other attached species; in the absence of the predator, one prey species
was capable of competitively excluding others. This keystone effect de-
pends upon two factors—that the prey assemblage structure is determined
by competition and that the predator preferentially feeds on the species
that is the best competitor in the assemblage. Clearly, not all food webs are
likely to be structured in this way.
Another method of experimental manipulation is to enhance the num-
bers of predators in a community. For complex natural communities, both
additions and exclusions of predators have revealed direct (depression of
prey) and indirect (enhancement) effects. Since generalist predators are
bitrophic in nature, they may interact with other carnivores in such a way
as to enhance the survival of herbivores that normally would fall victim. In
one experiment, adding praying mantises to an insect community resulted
in a decrease in spiders and a consequent increase in aphids, normally
eaten by these spiders. Such results are not uncommon and contribute to
the uncertainty of prediction. Mathematical models have been constructed
to depict predator-prey interactions in terms of how each population af-
fects the growth of the other. The simplest of these models, known as the
Lotka-Volterra model for the mathematicians who developed it, describes
a situation in which prey and predator populations are assumed to be mu-
tually regulating. This model, which was developed for a single prey pop-
ulation and single predator species, has been modified by many workers to
provide more realism, but it is far from predicting many competitive situa-
tions in complex natural communities.
As with the rest of modern ecology, these different approaches must be
blended in order to build a robust picture of how important predators are
in natural ecosystems. This knowledge would allow more successful pre-
diction of the outcomes of human intervention and more intelligent man-
agement of exploited populations. Predation is a key interaction in natural
ecosystems; understanding the nature of this interaction is central to any
understanding of nature itself.
Lawrence E. Hurd
541
Predation
See also: Balance of nature; Camouflage; Defense mechanisms; Displays;

Ethology; Food chains and webs; Hierarchies; Mammalian social systems;
Mimicry; Omnivores; Pheromones; Poisonous animals; Territoriality and
aggression.

Crawley, M. J., ed. Natural Enemies: The Population Biology of Predators, Para-
sites, and Diseases. Boston: Blackwell Scientific Publications, 1992.
Ehrlich, Paul R., and Anne H. Ehrlich. Extinction: The Causes and Conse-
quences of the Disappearance of Species. New York: Random House, 1981.
Fabre, Jean Henri. The Life of the Spider. Translated by Alexander Teixeira de
Mattos. New York: Dodd, Mead, 1916.
Gause, G. F. The Struggle for Existence. Reprint. New York: Dover, 1971.
Hassell, Michael P. Arthropod Predator-Prey Systems. Princeton, N.J.: Prince-
Krebs, J. R., and N. B. Davies, eds. Behavioral Ecology: An Evolutionary Ap-
proach. 4th ed. Boston: Blackwell Scientific Publications, 1997.
Levy, Charles Kingston. Evolutionary Wars: A Three-Billion Years Arms Race.
New York: Basingstoke, 2000.
Mowat, Farley. Never Cry Wolf. Boston: Atlantic-Little, Brown, 1963.
542
PUNCTUATED EQUILIBRIUM
VS. GRADUALISM
According to classical evolutionary theory, new species arise by gradual transfor-

mation of ancestral ones. Speciation theory of the 1950’s and 1960’s, however, pre-
dicted that new species arise from small populations isolated from the main popu-
lation, where they diverge rapidly.
I n 1972, Niles Eldredge and Stephen Jay Gould applied a new concept of
speciation to the fossil record, predicting that species should arise sud-
denly (“punctuated” by a speciation event) rather than gradually, and then
persist virtually unchanged for millions of years in “equilibrium” before
becoming extinct or speciating again.
Although Charles Darwin’s most influential work was entitled On the
Origin of Species by Means of Natural Selection (1859), in fact it did not ad-
dress the problem in the title. Darwin was concerned with showing that
evolution had occurred and that species could change, but he did not deal
with the problem of how new species were formed. For nearly a century,
no other biologists addressed this problem either. Darwin (and many of his
successors) believed that species formed by gradual transformation of ex-
isting ancestral species, and this viewpoint (known as gradualism) was
deeply entrenched in the biology and paleontology books for a century. In
this view, species are not real entities but merely arbitrary segments of con-
tinuously evolving lineages that are always in the process of change
through time. Paleontologists tried to document examples of this kind of
gradual evolution in fossils, but remarkably few examples were found.
The Allopatric Speciation Model

By the 1950’s and 1960’s, however, systematists (led by Ernst Mayr) began
to study species in the wild and therefore saw them in a different light.
They noticed that most species do not gradually transform into new ones
in the wild but instead have fairly sharp boundaries. These limits are estab-
lished by their ability and willingness to interbreed with each other. Those
individuals that can interbreed are members of the same species, and those
that cannot are of different species. When a population is divided and sep-
arated so that formerly interbreeding individuals develop differences that
prevent interbreeding, then a new species is formed. Mayr showed that, in
543
Punctuated equilibrium vs. gradualism
nature, large populations of individuals living together (sympatric condi-

tions) interbreed freely, so that evolutionary novelties are swamped out
and new species cannot arise.
When a large population becomes split by some sort of barrier so that
there are two different populations (allopatric conditions), however, the
smaller populations become isolated from interbreeding with the main
population. If these allopatric, isolated populations have some sort of un-
usual gene, their numbers may be small enough that this gene can spread
through the whole population in a few generations, giving rise to a new
species. Then, when the isolated population is reintroduced to the main
population, it has developed a barrier to interbreeding, and a new species
becomes established. This concept is known as the allopatric speciation
model.
The allopatric speciation model was well known and accepted by most
biologists by the 1960’s. It predicted that species arise in a few generations
from small populations on the fringe of the range of the species, not in the
main body of the population. It also predicted that the new species, once it
arises on the periphery, will appear suddenly in the main area as a new
species in competition with its ancestor. These models of speciation also
treated species as real entities, which recognize one another in nature and
are stable over long periods of time once they become established. Yet,
these ideas did not penetrate the thought of paleontologists for more than a
decade after biologists had accepted them.
Eldredge and Gould’s Model

In 1972, Niles Eldredge and Stephen Jay Gould proposed that the allopatric
speciation model would make very different predictions about species in
the fossil record than the prevailing dogma that they must change gradu-
ally and continuously through time. In their paper, they described a model
of “punctuated equilibrium.” Species should arise suddenly in the fossil
record (punctuation), followed by long periods of no change (equilibrium,
or stasis) until they went extinct or speciated again. They challenged pale-
ontologists to examine their biases about the fossil record and to see if in
fact most fossils evolved gradually or rapidly, followed by long periods of
stasis.
In the years since that paper, hundreds of studies have been done on
many different groups of fossil organisms. Although some of the data were
inadequate to test the hypotheses, many good studies have shown quite
clearly that punctuated equilibrium describes the evolution of many mul-
ticellular organisms. The few exceptions are in the gradual evolution of
size (which was specifically exempted by Eldredge and Gould) and in uni-
544
cellular organisms, which have both sexual and asexual modes of repro-
duction. Many of the classic studies of gradualism in oysters, heart ur-
chins, horses, and even humans have even been shown to support a model
of stasis punctuated by rapid change. The model is still controversial, how-
ever, and there are still many who dispute both the model and the data that
support it.
Implications
One of the more surprising implications of the model is that long pe-
riods of stasis are not predicted by classical evolutionary theory. In neo-
Darwinian theory, species are highly flexible, capable of changing in re-
sponse to environmental changes. Yet, the fossil record clearly shows that
most species persist unchanged for millions of years, even when other evi-
dence clearly shows climatic changes taking place. Instead of passively
changing in response to the environment, most species stubbornly persist
unchanged until they either go extinct, disappear locally, or change rapidly
to some new species. They are not infinitely flexible, and no adequate
mechanism has yet been proposed to explain the ability of species to main-
tain themselves in homeostasis in spite of environmental changes and
apparent strong natural selection. Naturally, this idea intrigues paleontol-
ogists, since it suggests processes that can only be observed in the fossil
record and were not predicted from studies of living organisms.
Species Selection
The punctuated equilibrium model has led to even more interesting ideas.
If species are real, stable entities that form by speciation events and split
into multiple lineages, then multiple species will be formed and compete
with one another. Perhaps some species have properties (such as the ability
to speciate rapidly, disperse widely, or survive extinction events) that give
them advantages over other species. In this case, there might be competi-
tion and selection between species, which was called species selection by
Steven Stanley in 1975. Some evolutionary biologists are convinced that
species selection is a fundamentally different process from that of simple
natural selection that operates on individuals. In species selection, the fun-
damental unit is the species; in natural selection, the fundamental unit is
the individual. In species selection, new diversity is created by speciation
and pruned by extinction; in natural selection, new diversity is created by
mutation and eliminated by death of individuals. There are many other
such parallels, but many evolutionary biologists believe that the processes
are distinct. Indeed, since species are composed of populations of individ-
uals, species selection operates on a higher level than natural selection.
545
If species selection is a valid description of processes occurring in na-

ture, then it may be one of the most important elements of evolution. Most
evolutionary studies in the past have concentrated on small-scale, or
microevolutionary, change, such as the gradual, minute changes in fruit
flies or bacteria after generations of breeding. Many evolutionary biolo-
gists are convinced, however, that microevolutionary processes are insuffi-
cient to explain the large-scale, or macroevolutionary, processes in the evo-
lution of entirely new body plans, such as birds evolving from dinosaurs.
In other words, traditional neo-Darwinism says that all evolution is merely
microevolution on a larger scale, whereas some evolutionary biologists
consider some changes too large for microevolution. They require differ-
ent kinds of processes for macroevolution to take place. If there is a differ-
ence between natural selection (a microevolutionary process) and species
selection (a macroevolutionary process), then species selection might be
a mechanism for the large-scale changes in the earth’s history, such as
great adaptive radiations or mass extinctions. Naturally, such radical ideas
are still controversial, but they are taken seriously by a growing number
of paleontologists and evolutionary biologists. If they are supported by
further research, then there may be some radical changes in evolutionary
biology.
Reinterpreting the Fossil Record

Determining patterns of evolution requires a very careful, detailed study
of the fossil record. To establish whether organisms evolve in a punctuated
or gradual mode, many criteria must be met. The taxonomy of the fossils
must be well understood, and there must be large enough samples at many
successive stratigraphic levels. To estimate the time spanned by the study,
there must be some form of dating that allows the numerical age of each
sample to be estimated. It is also important to have multiple sequences
of these fossils in a number of different areas to rule out the effects of mi-
gration of different animals across a given study area. Once the appropri-
ate samples have been selected, then the investigator should measure as
many different features as possible. Too many studies in the past have
looked at only one feature and therefore established very little. In particu-
lar, changes in size alone are not sufficient to establish gradualism, since
these phenomena can be explained by many other means. Finally, many
studies in the past have failed because they picked one particular lineage
or group and selectively ignored all the rest of the fossils in a given area.
The question is no longer whether one or more cases of gradualism or
punctuation occurs (they both do) but which is predominant among all the
organisms in a given study area. Thus, the best studies look at the entire as-
546
semblage of fossils in a given area over a long stratigraphic interval before

they try to answer the question of which tempo and mode of evolution is
prevalent.
Since the 1940’s, evolutionary biology has been dominated by the neo-
Darwinian synthesis of genetics, systematics, and paleontology. In more
recent years, many of the accepted neo-Darwinian mechanisms of evolu-
tion have been challenged from many sides. Punctuated equilibrium and
species selection represent the challenge of the fossil record to neo-Darwin-
ian gradualism and overemphasis on the power of natural selection. If fos-
sils show rapid change and long-term stasis over millions of years, then
there is no currently understood evolutionary mechanism for this sort of
stability in the face of environmental selection. A more general theory of
evolution may be called for, and, in more recent years, paleontologists, mo-
lecular biologists, and systematists have all been indicating that such a
radical rethinking of evolutionary biology is on the way.
Donald R. Prothero

tions and evolutionary explosions; Gene flow; Genetic drift; Genetically
modified foods; Isolating mechanisms; Natural selection; Nonrandom
mating, genetic drift, and mutation; Paleoecology; Population genetics;
Speciation; Species loss.

Bennett, K. D. Evolution and Ecology: The Pace of Life. New York: Cambridge
Eldredge, Niles. Time Frames: The Rethinking of Darwinian Evolution and the
Theory of Punctuated Equilibria. New York: Simon & Schuster, 1985.
Eldredge, N., and S. J. Gould. “Punctuated Equilibria: An Alternative to
Phyletic Gradualism.” In Models in Paleobiology, edited by T. J. M.
Schopf. San Francisco: Freeman, Cooper, 1972.
Freeman, S., and J. C. Herron. Evolutionary Analysis. 2d ed. Upper Saddle
Gerhart, John. Cells, Embryos, and Evolution: Toward a Cellular and Develop-
mental Understanding of Phenotypic Variation and Evolutionary Adaptabil-
ity. Malden, Mass.: Blackwell Science, 1997.
Gould, Stephen J. “The Meaning of Punctuated Equilibria and Its Role in
Validating a Hierarchical Approach to Macroevolution.” In Perspectives
547
on Evolution, edited by Roger Milkman. Sunderland, Mass.: Sinauer As-

sociates, 1982.
Gould, Stephen J., and Niles Eldredge. “Punctuated Equilibrium: The
Tempo and Mode of Evolution Reconsidered.” Paleobiology 3 (1977):
115-151.
Hoffman, Antoni. Arguments on Evolution: A Paleontologist’s Perspective.
New York: Oxford University Press, 1988.
Levinton, Jeffrey S. Genetics, Paleontology, and Macroevolution. 2d ed. New
Mayr, Ernst. Animal Species and Evolution. Cambridge, Mass.: Harvard Uni-
Moller, A. P. Asymmetry, Developmental Stability, and Evolution. New York:
Oxford University Press, 1997.
548
RAIN FORESTS
A forest growing in a region that receives over one hundred inches of rain annually
is considered to be a rain forest. Rain forests can be found in both tropical and tem-
perate climates and are noted for their remarkable biodiversity. Thousands of dif-
ferent animal and plant species can be found within only an acre or two of a rain
forest.
R ain forests are forests found in regions of the world that receive large
amounts of precipitation annually. Rain forests present an incredibly
diverse range of habitats, as they exist both at low elevations and high in
mountain ranges. Many unusual and seldom-seen creatures inhabit the
world’s rain forests, including spiders so large they eat small birds, and
colorful but highly poisonous tree frogs. The enigmatic sloth, an animal
that spends its entire life hanging upside down from tree limbs and mov-
ing so slowly that moss grows on its fur, is found in the rain forests of South
America.
Although tropical rain forests, such as those in the Amazon River drain-
age system of South America, are perhaps the best known, rain forests do
exist in temperate regions as well. Olympic National Park in the state of
Washington preserves a temperate climate rain forest, while much of the
coast of British Columbia and southeastern Alaska also receives well over
one hundred inches of rain annually. The primary difference between tem-
perate and tropical rain forests is that in a temperate rain forest, often one
or two species of trees will become dominant. In the coniferous rain forest
of the Pacific Northwest, for example, Douglas fir and western red cedar
are the dominant species, while other trees are found in much smaller
numbers. In a tropical rain forest, in contrast, several hundred species of
trees may grow side by side within a very small geographic area. The ma-
jority of trees found in tropical rain forests tend to be broad-leaved, such as
the rubber tree, while temperate rain forests are dominated by conifers.
The leaves of many plants in rain forests often have a waxy texture or come
to a point to help shed water more quickly and prevent the growth of fungi
or mold.
Characteristics of Rain Forests

Although rain forests are remarkably diverse, they do share a few charac-
teristics in common. The abundant moisture in a rain forest gives the
549
Rain forests
woodland a lush, fertile appearance. This is particularly true in tropical re-

gions. Even in the understory, close to ground level where light is limited,
vegetation may be dense. This appearance of fertility is often deceptive.
Dead plant matter decays rapidly in a tropical forest, but the nutrients are
used quickly by the numerous competing plants. In addition, the trees in
tropical forests are evergreen, which means the litter that does fall to the
forest floor does so irregularly, unlike temperate broadleaf forests, where
trees lose their leaves annually as the seasons change. Leaves will remain
indefinitely on tropical species, such as fig and rubber trees, which is one
reason small specimens of these trees are popular as houseplants. As a con-
sequence of this lack of mulch, topsoil is often thin and the root systems of
the trees are quite shallow.
One reason tropical rain forests are evergreen is that in the tropics there
is little seasonal variation in the hours of daylight. The closer to the equator
a forest lies, the less change there is from season to season. In temperate cli-
mates, many plants have evolved to bloom, set seeds, or lose their leaves
based on the number of hours of sunlight available each day. As the sea-
Although most think of rain

forests as tropical
ecosystems, many coastal or
near-coastal forests in
higher latitudes, such as
those of the Pacific
Northwest, fall into this
category as well, with their
high rates of precipitation
and their various levels of
growth. Here, understory
and canopy are clearly
visible. (PhotoDisc)
550
Rain forests
sons change annually, plants bloom in the spring or early summer; fruit
ripens in the fall, photosynthesis slows, and leaves change color and die. In
the tropics, where the number of hours of sun light daily never varies,
plants follow a different schedule. Many tropical plants bear new flowers
and mature fruit simultaneously. The evergreen foliage and continuous
supply of certain fruits has led to the adaptation of some animals to a very
restricted diet: koalas, for example, which feed exclusively on eucalyptus
leaves, or parakeets that eat only figs. Exceptions to this pattern are the for-
ests where rain fall is seasonal, such as regions of the world like southeast
Asia, where much of the rain comes in the form of annual monsoon storms.
In those cases, flowering and setting fruit will coincide with the seasonal
rains.
Forest Zones
A rain forest can be divided into four zones, each of which has its own dis-
tinct characteristics. The lowest level, the forest floor, is often dark and
gloomy. Little sunlight penetrates to this level, and there is little air move-
ment. Numerous insects, such as beetles, cockroaches, and termites, live in
the decaying litter and provide food for larger animals and birds. Many of
the insects, birds, reptiles, and amphibians that live in the lower levels of
the rain forest are brightly colored. Scientists speculate that the animals
have evolved in this fashion to more easily attract potential mates. Other
scientists believe that colors warn potential predators to stay away. In ei-
ther case, the vivid colors make the animals more easily seen in what is oth-
erwise a dark environment.
Just above the forest floor is the understory. Many of the plants in the
understory have large, dark leaves to maximize their light-collecting abil-
ity. Because there is little natural air movement within the lower levels of a
rain forest due to the canopy blocking any natural breezes, the flowering
plants in the understory often have strongly scented or vividly colored
flowers to help attract insects or birds to assist with pollination. Lizards,
snakes, amphibians such as tree frogs and salamanders, small birds, and
mammals as large as the jaguar all call the understory home. The plants
found only in the understory seldom exceed fifteen to twenty feet in
height. The coffee shrub is an example of a small, shade-tolerant, tropical
tree. Until horticulturalists developed strains of coffee for use in planta-
tions where the coffee bushes are the only plants grown, coffee grew natu-
rally in the understory of tropical forests.
The densest layer of plant life is the canopy. High above the rain forest
floor, the branches of mature trees form a dense intertwined zone of vege-
tation extending up as much as 150 feet above the ground. Numerous
551
Rain forests
plants sprout in the crotches of trees, where debris may collect. Tree limbs
are festooned with vines and mosses, and bromeliads and orchids grow on
the rough bark of tree trunks. Even other trees may start their life cycle a
hundred feet above the ground: The strangler figs of Borneo are a relatively
shade-intolerant species. A fig seed that lands and sprouts on the ground
will probably not survive due to low light levels on the forest floor. Stran-
gler figs have adapted so that their seedlings do best high in the canopy.
The figs begin life in the crotches of other trees. The roots of a young fig will
gradually creep down the trunk of the tree on which it sprouted. Over
time, the strangler fig’s roots will completely encircle the host tree and pen-
etrate the forest floor. The fig thrives, but the host tree dies, choked off by
the strangler fig. Primates such as gibbons, orangutans, and lemurs spend
much of their lives in the canopy, feeding on the fruit of trees such as the
strangler fig, as do the sloth and other herbivorous mammals.
The emergent layer of the rain forest consists of the tallest trees, some of
which exceed two hundred feet in height. The tops of these trees provide a
habitat for large, predatory birds, such as eagles, as well as being home to
assorted snakes, monkeys, and other animals. Every layer of the rain forest
teems with life, and often what can be found at ground level gives no hint
of the diversity that exists two hundred feet above in the tree tops.
Rain Forest Conservation

Many of the trees found in rain forests are valued for their commercial use
as lumber, while others have been exploited for their fruits or other prod-
ucts, causing much habitat loss. Tropical hardwoods, such as teak and ma-
hogany, for example, have long been used in construction and in furniture.
Teak resists rotting and as a result is often used for products that are going
to be exposed to the weather, such as garden furniture. Because teak is de-
sirable as lumber, timber companies are increasingly planting it in planta-
tions for a sustainable yield rather than relying solely on natural forests as
a source.
Activists hoping to preserve the tropical rain forest have encouraged in-
digenous peoples to collect forest products, such as nuts or sap, as a way to
create a viable economy while at the same time discouraging industrial
clear-cutting of the forest. Native people tap rubber trees in Amazonia, for
example, to collect latex. Rubber trees are native to the rain forests of South
America, although they are also grown in plantations in other tropical re-
gions of the world, such as Southeast Asia.
The biggest threat to the world’s rain forests may not come from com-
mercial logging, however. In many regions of the world, rain forests have
fallen victim to population pressures. Forests continue to be clear-cut for
552
Rain forests
agricultural use, even when the farmers and ranchers know the exposed
soil’s fertility will be quickly exhausted. In some cases, the cleared land be-
comes an arid wasteland as the tropical sun bakes the soil too hard to ab-
sorb rain water. In others, the land is farmed for a year or two and then
abandoned. Given enough time, the rain forest may regenerate, but the
process will take hundreds of years.

ests; Grasslands and prairies; Habitats and biomes; Lakes and limnology;
Marine biomes; Mediterranean scrub; Mountain ecosystems; Old-growth
forests; Rain forests and the atmosphere; Rangeland; Reefs; Savannas and
deciduous tropical forests; Taiga; Tundra and high-altitude biomes; Wet-
lands.

Bowman, David M. J. S. Australian Rainforests: Islands of Green in a Land of
Fire. New York: Cambridge University Press, 2000.
Durbin, Kathie. Pulp Politics and the Fight for the Alaska Rain Forest. Cor-
vallis: Oregon State University Press, 1999.
Gamlin, Linda, and Anuschka de Rohan. Mysteries of the Rain Forest.
Pleasantville, N.Y.: Reader’s Digest, 1998.
Goldsmith, Frank Barrie. Tropical Rain Forest: A Wider Perspective. New
York: Chapman & Hall, 1998.
Holloway, M. “Sustaining the Amazon.” Scientific American 269 (July, 1993):
90-99.
Killman, Wolf, and Lay Thong Hong. “Rubberwood: The Success of an Ag-
ricultural By-Product.” Unasylva 51 (2000): 66-72.
Maser, Chris. Forest Primeval: The Natural History of an Ancient Forest. Re-
print. Corvallis: Oregon State University Press, 2001.
Tricart, Jean. The Landforms of the Humid Tropics, Forests, and Savannas.
Translated by Conrad J. Kiewiet de Jonge. New York: St. Martin’s Press,
1972.
553
RAIN FORESTS AND THE
ATMOSPHERE
Types of ecology: Biomes; Ecoenergetics; Ecosystem ecology; Global

ecology
Because photosynthesis releases large amounts of oxygen into the air, a curtail-
ment of the process by rain-forest deforestation may have negative effects on the
global atmosphere.
R ain forests are ecosystems noted for their high biodiversity and high
rate of photosynthesis. The rapid deforestation of such areas is of
great concern to environmentalists both because it may lead to the extinc-
tion of numerous species and because it may reduce the amount of photo-
synthesis occurring on the earth.
All living things on the earth—plants, animals, and microorganisms—
depend on the “sea” of air surrounding them. The atmosphere includes
abundant, permanent gases such as nitrogen (78 percent) and oxygen (21
percent) as well as smaller, variable amounts of other gases such as water
vapor and carbon dioxide. Organisms absorb and use this air as a source of
raw materials and release into it by-products of their life activities.
Cellular Respiration
Cellular respiration is the most universal of the life processes. A series of
chemical reactions beginning with glucose and occurring in cytoplasmic
organelles called mitochondria, cellular respiration produces a chemical
compound called adenosine triphosphate (ATP). This essential substance
furnishes the energy cells need to move, to divide, and to synthesize chem-
ical compounds—in essence, to perform all the activities necessary to sus-
tain life. Cellular respiration occurs in plants as well as animals, and it oc-
curs during both the day and the night. In order for the last of the series of
chemical reactions in the process to be completed, oxygen from the sur-
rounding air (or water, in the case of aquatic plants) must be absorbed. The
carbon dioxide that forms is released into the air.
For cellular respiration to occur, a supply of glucose (a simple carbohy-
drate compound) is required. Photosynthesis, an elaborate series of chemi-
cal reactions occurring in chloroplasts, produces glucose, an organic car-
bon compound with six carbon atoms. Energy present in light must be
trapped by the chlorophyll within the chloroplasts to drive photosynthe-
554
Rain forests and the atmosphere
sis. Therefore, photosynthesis occurs only in plants and related organisms

such as algae, and only during the daytime. Carbon dioxide, required as a
raw material, is absorbed from the air, while the resulting oxygen is re-
leased into the atmosphere. The exchange of gases typically involves tiny
openings in leaves, called stomata.
Oxygen Cycle
Oxygen is required for the survival of the majority of microorganisms and
all plants and animals. From the surrounding air, organisms obtain the ox-
ygen used in cell respiration. Plants absorb oxygen through the epidermal
coverings of their roots and stems and through the stomatal openings of
their leaves.
The huge amounts of oxygen removed from the air during respiration
must be replaced in order to maintain a constant reservoir of oxygen in the
atmosphere. There are two significant sources of oxygen. One involves
water molecules of the atmosphere that undergo a process called photo-
dissociation: Oxygen remains after the lighter hydrogen atoms are re-
leased from the molecule and escape into outer space.
An aerial view of the

rain forest in Guyana. In
such tropical forests, the
many layers of forest
vegetation result in
energy from sunlight
being efficiently used as
it passes downward. The
large amounts of oxygen
released are available for
use not only by the
forests but also, because
of global air movement,
by other ecosystems
throughout the world.
(PhotoDisc)
555
The other source is photosynthesis. Chlorophyll-containing organisms

release oxygen as they use light as the energy source to split water mole-
cules in a process called photolysis. The hydrogen is transported to the ter-
minal phase of photosynthesis called the Calvin cycle, where it is used as
the hydrogen source necessary to produce and release molecules of the car-
bohydrate glucose. In the meantime, the oxygen from the split water is re-
leased into the surrounding air.
Early in the history of the earth, before certain organisms evolved the
cellular machinery necessary for photosynthesis, the amount of atmo-
spheric oxygen was very low. As the number and sizes of photosynthetic
organisms gradually increased, so did the levels of oxygen in the air. A pla-
teau was reached several million years ago as the rate of oxygen release
and absorption reached an equilibrium.
Ozone
Another form of oxygen is ozone. Unlike ordinary atmospheric oxygen, in
which each molecule contains two atoms, ozone molecules have three oxy-
gen atoms each. Most ozone is found in the stratosphere at elevations be-
tween 10 and 50 kilometers (6 and 31 miles). This layer of ozone helps to
protect life on earth from the harmful effects of ultraviolet radiation. Scien-
tists, especially ecologists, are concerned because the amount of ozone has
been reduced drastically over the last few decades. Already, an increase in
the incidence of skin cancer in humans and a decrease in the efficiency of
photosynthesis has been documented. Another concern related to ozone is
that of an increase in ozone levels nearer to the ground, where living things
are harmed as a result. The formation of ozone from ordinary oxygen
within the atmosphere is greatly accelerated by the presence of gaseous
pollutants released from industrial processes.
Carbon Cycle
All forms of life are composed of organic (carbon-containing) molecules.
Carbohydrates include glucose as well as lipids (fats, oils, steroids, and
waxes), proteins, and nucleic acids. The ability of carbon to serve as the
backbone of these molecules results from the ability of carbon atoms to
form chemical bonds with other carbon atoms and also with oxygen, hy-
drogen, and nitrogen atoms.
Like oxygen, carbon cycles in a predictable manner between living
things and the atmosphere. In photosynthesis, carbon is “fixed” as carbon
dioxide in the air (or dissolved in water) is absorbed and converted into
carbohydrates. Carbon cycles to animals as they feed on plants and algae.
As both green and nongreen organisms respire, some of their carbohy-
556
drates are oxidized, releasing carbon dioxide into the air. Each organism
must eventually die, after which decay processes return the remainder of
the carbon to the atmosphere.
Greenhouse Effect
Levels of atmospheric carbon dioxide have fluctuated gradually during
past millennia, as revealed by the analysis of the gas trapped in air bubbles
of ice from deep within the earth. In general, levels were lower during gla-
cial periods and higher during warmer ones. After the nineteenth century,
levels rose slowly until about 1950 and then much more rapidly afterward.
The apparent cause has been the burning of increased amounts of fossil fu-
els associated with the Industrial Revolution and growing energy de-
mands in its wake. The global warming that is now being experienced is
believed by most scientists to be the cause of increased carbon dioxide lev-
els. The greenhouse effect is the term given to the insulating effects of the
atmosphere with increased amounts of carbon dioxide. The earth’s heat is
lost to outer space less rapidly, thus increasing the earth’s average temper-
ature.
Forest Ecosystems
The biotic (living) portions of all ecosystems include three ecological or
functional categories: producers (plants and algae), consumers (animals),
and decomposers (bacteria and fungi). The everyday activities of all organ-
isms involve the constant exchange of oxygen and carbon dioxide between
the organisms of all categories and the surrounding atmosphere.
Because they release huge quantities of oxygen during the day, produc-
ers deserve special attention. In both fresh and salt water, algae are the
principal producers. On land, this role is played by a variety of grasses,
other small plants, and trees. Forest ecosystems, dominated by trees but
also harboring many other plants, are major systems that produce a dis-
proportionate amount of the oxygen released into the atmosphere by ter-
restrial ecosystems.
Forests occupy all continents except for Antarctica. A common classifi-
cation of forests recognizes these principal categories: coniferous (northern
evergreen), temperate deciduous, and tropical evergreen, with many sub-
categories for each. The designation “rain forest” refers to the subcatego-
ries of these types that receive an amount of rainfall well above the aver-
age. Included are tropical rain forests (the more widespread type) and
temperate rain forests. Because of the ample moisture they receive, both
types contain lush vegetation that produces and releases oxygen into the
atmosphere on a larger scale than do other forests.
557
Tropical Rain Forests

Tropical rain forests exist at relatively low elevations in a band about the
equator. The Amazon basin of South America contains the largest continu-
ous tropical rain forest. Other large expanses are located in western and
central Africa and the region from Southeast Asia to Australia. Smaller ar-
eas of tropical rain forests occur in Central America and on certain islands
of the Caribbean Sea, the Pacific Ocean, and the Indian Ocean. Seasonal
changes within tropical rain forests are minimal. Temperatures, with a
mean near 25 degrees Celsius, seldom vary more than 4 degrees Celsius.
Rainfall each year measures at least 400 centimeters.
Tropical rain forests have the highest biodiversity of any terrestrial eco-
system. Included is a large number of species of flowering plants, insects,
and animals. The plants are arranged into layers, or strata. In fact, all for-
ests are stratified but not to the same degree as tropical rain forests. A ma-
ture tropical rain forest typically has five layers. Beginning with the upper-
most, they are an emergent layer (the tallest trees that project above the
next layer); a canopy of tall trees; understory trees; shrubs, tall herbs, and
ferns; and low plants on the forest floor.
Several special life-forms are characteristic of the plants of tropical rain
forests. Epiphytes are plants such as orchids that are perched high in the
branches of trees. Vines called lianas wrap themselves around trees. Most
tall trees have trunks that are flared at their bases to form buttresses that
help support them in the thin soil.
This brief description of tropical rain forests helps to explain their role
in world photosynthesis and the related release of oxygen into the atmo-
sphere. As a result of the many layers of forest vegetation, the energy from
sunlight as it passes downward is efficiently utilized. Furthermore, the
huge amounts of oxygen released are available for use not only by the for-
ests themselves but also, because of global air movement, by other ecosys-
tems throughout the world. Because of this, tropical rain forests are often
referred to as “the earth’s lungs.”
Temperate Rain Forests

Temperate rain forests are much less extensive than tropical rain forests;
they occur primarily along the Pacific Coast in a narrow band from south-
ern Alaska to central California. Growing in this region is a coniferous for-
est but one with warmer temperatures and a higher rainfall than those to
the north and inland. This rainfall of 65 to 400 centimeters per year is much
less than that of a tropical rain forest, but is supplemented in the summer
by frequent heavy fogs. As a result, evaporation rates are greatly reduced.
Because of generally favorable climatic conditions, temperate rain forests,
558
like tropical ones, support a lush vegetation. The rate of photosynthesis

and release of oxygen are higher than in most other world ecosystems.
Ecologists and conservationists are greatly concerned about the mas-
sive destruction of rain forests. Rain forests are being cut and burned at a
rapid rate to plant crops, to graze animals, and to provide timber. The ulti-
mate effect of deforestation of these special ecosystems is yet to be seen.
Thomas E. Hemmerly
See also: Biodiversity; Biomes: determinants; Biomes: types; Commu-

nities: ecosystem interactions; Deforestation; Ecosystems: definition and
history; Ecosystems: studies; Forests; Geochemical cycles; Global warm-
ing; Greenhouse effect; Habitats and biomes; Hydrologic cycle; Nutrient
cycles; Ozone depletion and ozone holes; Pollution effects; Rain forests; Sa-
vannas and deciduous tropical forests; Slash-and-burn agriculture; Trophic
levels and ecological niches.

Laurance, William F., and Richard O. Bierregaard, eds. Tropical Forest Rem-
nants: Ecology, Management, and Conservation of Fragmented Communities.
Chicago: University of Chicago Press, 1997.
Shipp, Steve. Rainforest Organizations: A Worldwide Directory of Private and
Governmental Entities. Jefferson, N.C.: McFarland, 1997.
Townsend, Janet G. Women’s Voices from the Rainforest. New York: Rout-
ledge, 1995.
Vandermeer, John. Breakfast of Biodiversity: The Truth About Rain Forest De-
struction. Oakland, Calif.: Institute for Food and Development Policy,
1995.
559
RANGELAND
Types of ecology: Agricultural ecology; Biomes; Ecosystem ecology
Open land of a wide variety of types, including grasslands, shrublands, marshes,

and meadows as well as some desert and alpine land, is known as rangeland.
R angeland is a valuable and resilient ecosystem resource that supports

considerable plant and animal life. Rangeland generally refers to a
kind of land rather than a use of that land. The Society for Range Manage-
ment defines rangelands as “land on which the native vegetation (climax
or natural potential) is predominantly grasses, grass-like plants, forbs, or
shrubs.” Rangeland “includes lands revegetated naturally or artificially”
as well as “natural grasslands, savannas, shrublands, most deserts, tundra,
alpine communities, coastal marshes and wet meadows.”
Rangelands usually have some limitation for intensive agriculture,
such as low and erratic precipitation, lack of soil fertility, shallow or rocky
soil, or steep slopes. In addition to livestock grazing, rangelands serve
multiple-use functions such as providing recreational opportunities, wa-
tersheds, mining locations, and habitat for many animal species. Renew-
able natural resources associated with rangelands are plants and animals
(and, in some senses, water). Nonrenewable resources include minerals
and other extractable materials.
Location and Characteristics

Rangelands are extensive and extremely variable. As defined by the Soci-
ety for Range Management, they occupy more than 50 percent of the
world’s total land surface and about 1 billion acres in the United States
alone. Rangelands are home to nomadic herders on nearly every continent.
They vary from high-elevation alpine tundra and high-latitude Arctic tun-
dra to tropical grasslands. The tall-grass prairies in the United States (now
mostly plowed for intensive agriculture) and the rich grasslands of eastern
Africa are among the most productive.
Rangelands grade into woodlands and forests as woody species and
trees become more abundant. Some forests are grazed by wild and domes-
tic animals, and the distinction between rangeland and forest is often not
clear. The other difficult distinction is between rangeland and pastureland.
Pastureland is generally improved by seeding, fertilization, or irrigation,
whereas rangelands support native plants and have little intensive im-
provement.
560
Rangeland
In the United States, rangeland improvements during the twenty years

following World War II often included brush control, grazing manage-
ment, and seeding, but rangelands were not irrigated. After the 1970’s,
when fuel costs increased and environmental concerns about pesticide use
increased, brush control practices were reduced considerably. Today envi-
ronmental concerns include rangeland degradation from overgrazing, es-
pecially on riparian vegetation along streams, and concern for endangered
animal and plant species. These issues have become controversial in the
United States.
Rangelands as Ecosystems
Rangelands constitute natural ecosystems with nonliving environmental
factors such as soil and climatic factors. Life-forms are primary producers
(grasses, forbs, and shrubs), herbivores (livestock; big game animals such
as deer and bison; and many rodents and insects), carnivores (such as coy-
otes, bears, and eagles), and decomposers (fungi and bacteria) that break
down organic matter into elements that can be utilized by plants. Plants
convert carbon dioxide and water into complex carbohydrates, fats, and
proteins that nourish animals feeding on the plants.
Image Not Available
561
Rangeland
Individual chemical elements are circulated throughout the various

components. Many of these elements are present in the soil, including
phosphorus, magnesium, potassium, and sulfur. Nitrogen, on the other
hand, is present in large amounts in the atmosphere but must be converted
(fixed) into forms that can be utilized by plants before it can be cycled. En-
ergy is fixed through the process of photosynthesis and transformed to
forms useful for the plants, then the animals that feed on plants.
When chemicals are taken up by plant roots from the soil, they become
available to a wide group of herbivores, from small microbes to large
ungulates. Eventually nutrients are passed on to organisms at higher
trophic levels (omnivores and carnivores). Both plant and animal litter is
eventually broken down by decomposers—bacteria, fungi, and other soil
organisms—and returned to the soil or, in the case of carbon or nitrogen,
given off to the atmosphere.
However, energy is degraded at each step along the way; energy is
transferred but not cycled. Grazing animals on rangelands influence
plants by removing living tissue, by trampling, and by altering competi-
tive relations with other plants. Large grazing animals tend to compact the
soil, reducing infiltration and increasing surface runoff.
Rangeland Dynamics
Rangelands vary considerably with time. Scientists are gaining a better un-
derstanding of some factors related to rangeland change. Pollen records
and, in the southwestern United States, packrat middens have been used
to reconstruct past climate and vegetational conditions. Some areas have
become drier and others more mesic. The formation and retreat of glaciers
influenced climatic patterns and soil development. A recent general trend
in many rangelands is an increase in woody plants at the expense of
grasses. Many factors are probably responsible for these shifts, but fire con-
trol, overgrazing, climatic shifts, introduction of exotic species, and influ-
ence of native animals are likely causal agents.
Rangelands are being threatened by encroachment from crop agri-
culture as worldwide development increases. Nomadic herders tradi-
tionally met periodic drought conditions by having the flexibility to move
to areas not impacted by drought. Now, with area lost to livestock graz-
ing and other political restrictions, herders are often forced to maintain
higher livestock numbers to support those directly dependent on live-
stock. Despite various kinds of disturbances and stresses on rangelands,
these areas have supported many large grazing animals and people for
centuries.
Rex D. Pieper
562
Rangeland
See also: Forest management; Grasslands and prairies; Grazing and over-
grazing; Multiple-use approach.

Heady, Harold F., and R. Dennis Child. Rangeland Ecology and Management.
2d ed. Boulder, Colo.: Westview, 2000.
Holechek, Jerry L., Rex D. Pieper, and Carlton H. Herbel. Range Manage-
ment: Principles and Practices. 4th ed. Upper Saddle River, N.J.: Prentice
Hall, 2001.
Jacobs, Lynn. Waste of the West: Public Lands Ranching. Tucson, Ariz.:
L. Jacobs, 1991.
Longworth, John W., and Gregory J. Williamson. China’s Pastoral Region:
Sheep and Wool, Minority Nationalities, Rangeland Degradation, and Sus-
tainable Development. Wallingford, England: CAB International, 1993.
Owen, Oliver S., Daniel D. Chiras, and John P. Reganold. Natural Resource
Conservation. 7th ed. Upper Saddle River, N.J.: Prentice Hall, 1998.
Sayre, Nathan F. The New Ranch Handbook: A Guide to Restoring Western
Rangelands. Santa Fe, N.Mex.: Quivera Coalition, 2001.
563
REEFS
ecology
Reefs are among the oldest known communities, existing at least 2 billion years
ago. They exert considerable control on the surrounding physical environment, in-
fluencing turbulence levels and patterns of sedimentation. Ancient reefs are often
important hydrocarbon reservoirs.
“True” Reefs vs. Reeflike Structures

Reefs or reeflike structures are among the oldest known communities, ex-
tending back more than 2 billion years into the earth’s history. These earli-
est reefs were vastly different in their biotic composition and physical
structure from modern reefs, which are among the most diverse of biotic
communities and display amazingly high rates of biotic productivity (car-
bon fixation) and calcium carbonate deposition, despite their existence in a
virtual nutrient “desert.” Reefs are among the few communities to rival the
power of humankind as a shaper of the planet. The Great Barrier Reef of
Australia, for example, forms a structure some 2,000 kilometers in length
and up to 150 kilometers in width.
It is necessary to distinguish between “true,” or structural, reefs and
reeflike structures or banks. Reefs are carbonate structures that possess an
internal framework. The framework traps sediment and provides resis-
tance to wave action; thus, reefs can exist in very shallow water and may
grow to the surface of the oceans. Banks are also biogenically produced but
lack an internal framework. Thus, banks are often restricted to low-energy,
deep-water settings. “Bioherm” refers to moundlike carbonate buildups,
either reefs or banks, and “biostrome” to low, lens-shaped buildups.
Reef Classification
Modern reefs are classified into several geomorphic types: atoll, barrier,
fringing, and patch. Many of these may be further subdivided into reef
crest or flat, back-reef or lagoon, and fore-reef zones. Atoll reefs are circular
structures with a central lagoon, thought to form on subsiding volcanic is-
lands. Barrier reefs are elongate structures that parallel coastlines and pos-
sess a significant lagoon between the exposed reef crest and shore. These
often occur on the edges of shelves that are uplifted by faulting. Fringing
reefs are elongate structures paralleling and extending seaward from the
coastline that lack a lagoon between shore and exposed reef crest. Patch
564
Reefs
reefs are typically small, moundlike structures, occurring isolated on

shelves or in lagoons. The majority of fossil reefs would be classified as
patch reefs, although many examples of extensive, linear, shelf-edge trends
are also known from the geologic record.
Reefs form one of the most distinctive and easily recognized sedimen-
tary facies (or environments). In addition to possessing a characteristic
fauna consisting of corals, various algae, and stromatoporoids, they are
distinguished by a massive (nonlayered) core that has abrupt contacts with
adjacent facies. Associated facies include flat-lying lagoon and steeply in-
clined fore-reef talus, the latter often consisting of large angular blocks de-
rived from the core. The reef core is typically a thick unit relative to adja-
cent deposits. The core also consists of relatively pure calcium carbonate
with little contained terrigenous material.
Reef Environments
Modern reefs are restricted to certain environments. They occur abun-
dantly only between 23 degrees north and south latitudes and tend to
be restricted to the western side of ocean basins, which lack upwelling of
cold bottom waters. This restriction is based on temperature, as reefs do
not flourish where temperatures frequently reach below 18 degrees Cel-
sius. Reef growth is largely restricted to depths greater than 60 meters, as
there is insufficient penetration of sunlight below this depth for symbiont-
bearing corals to flourish. Reefs also require clear waters lacking sus-
pended terrigenous materials, as these interfere with the feeding activ-
ity of many reef organisms and also reduce the penetration of sunlight.
Finally, most reef organisms require salinities that are in the normal oce-
anic range. It appears that many fossil reefs were similarly limited in their
environmental requirements.
Some of the most striking features of modern reefs include their pro-
nounced zonation, great diversity, and high productivity and growth rates.
Reefs demonstrate a strong bathymetric (depth-related) zonation. This
zonation is largely mediated through depth-related changes in turbulence
intensity and in the quantity and spectral characteristics (reds are absorbed
first, blues last) of available light. Shallow (1- to 5-meter) fore-reef environ-
ments are characterized by strong turbulence and high light intensity and
possess low-diversity assemblages of wave-resistant corals, such as the
elk-horn coral, Acropora palmata, and crustose red algae.
With increasing depth (10-20 meters), turbulence levels decrease and
coral species diversity increases, with mound and delicate branching colo-
nies occurring. At greater depths (30-60 meters), corals assume a flattened,
platelike form in an attempt to maximize surface area for exposure to am-
565
Reefs
bient light. Sponges and many green algae are also very important over
this range. Finally, corals possessing zooxanthellae, which live in the coral
tissues and provide food for the coral host, are rare or absent below 60 me-
ters because of insufficient light. Surprisingly, green and red calcareous al-
gae extend to much greater depths (100-200 meters), despite the very low
light intensity (much less than 1 percent of surface irradiance). Sponges are
also important members of these deep reef communities.
Diveristy of Life-Forms
Coral reefs are among the most diverse of the earth’s communities; how-
ever, there is no consensus on the mechanism(s) behind the maintenance of
this great diversity. At one time, it was believed that reefs existed in a low-
disturbance, highly stable environment, which allowed very fine subdivi-
sion of food and habitat resources and thus permitted the coexistence of a
great number of different species. Upon closer inspection, however, many
reef organisms appear to overlap greatly in food and habitat requirements.
Also, it has become increasingly apparent that disturbance, in the form of
disease, extreme temperatures, and hurricanes, is no stranger to reef com-
munities.
Coral reefs exhibit very high rates of productivity (carbon fixation),
which is a result of extremely tight recycling of existing nutrients. This is
necessary, as coral reefs exist in virtual nutrient “deserts.” Modern corals
exhibit high skeletal growth rates, up to 10 centimeters per year for some
branching species. Such high rates of skeletal production are intimately re-
lated to the symbiosis existing between the hermatypic or reef-building
scleractinian corals (also gorgonians and many sponges) and unicellular
algae or zooxanthellae. Corals that, for some reason, have lost their zo-
oxanthellae or that are kept in dark rooms exhibit greatly reduced rates of
skeleton production.
In addition to high individual growth rates for component taxa, the car-
bonate mass of the reefs may grow at a rate of some 2 meters per 1,000
years, a rate that is much higher than that of most other sedimentary de-
posits. This reflects the high productivity or growth rates of the component
organisms and the efficient trapping of derived sediment by the reef frame.
Although the framework organisms, most notably corals, are perhaps the
most striking components of the reef system, the framework represents
only 10-20 percent of most fossil reef masses. The remainder of the reef
mass consists of sedimentary fill derived from the reef community through
a combination of biosynthesis (secretion) and bioerosion (breaking down)
of calcium carbonate. An example of the relative contributions of reef or-
ganisms to sediment can be found in Jamaica, where shallow-water, back-
566
Reefs
This underwater close-up of a

portion of Australia’s Great
Barrier Reef displays the diversity
of species supported by this
marine ecosystem. The Great
Barrier Reef forms a structure
some 2,000 kilometers in length
and up to 150 kilometers in
width. (Corbis)
reef sediment consists of 41 percent coral, 24 percent green calcareous al-

gae, 13 percent red calcareous algae, 6 percent foraminifera, 4 percent mol-
lusks, and 12 percent other grains. The most important bioeroders are bor-
ing sponges, bivalves, and various “worms,” which excavate living spaces
within reef rock or skeletons, and parrot fish and sea urchins, which re-
move calcium carbonate as they feed upon surface films of algae.
Types of Reef Communities

A diversity of organisms has produced reef and reeflike structures through-
out the earth’s history. Several distinct reef community types have been
noted, as well as four major “collapses” of reef communities. The oldest
reefs or reeflike structures existed more than 2 billion years ago during the
Precambrian eon. These consisted of low-diversity communities domi-
nated by soft, blue-green algae, which trapped sediment to produce lay-
ered, often columnar structures known as stromatolites.
During the Early Cambrian period, blue-green algae were joined by cal-
careous, conical, spongelike organisms known as archaeocyathids, which
persisted until the end of the Middle Cambrian. Following the extinction
of the archaeocyathids, reefs again consisted only of blue-green algae until
the advent of more modern reef communities in the Middle Ordovician pe-
riod. These reefs consisted of corals (predominantly tabulate and, to a
much lesser extent, rugose corals), red calcareous algae, bryozoans (moss
567
Reefs
animals), and the spongelike stromatoporoids. This community type per-

sisted through the Devonian period, at which time a global collapse of reef
communities occurred.
The succeeding Carboniferous period largely lacked reefs, although al-
gal and crinoidal (sea lily) mounds were common. Reefs again occurred in
the Permian period, consisting mainly of red and green calcareous algae,
stromatolites, bryozoans, and chambered calcareous sponges known as
sphinctozoans, which resembled strings of beads. These reefs were very
different from those of the earlier Paleozoic era; in particular, the tabulates
and stromatoporoids no longer played an important role. The famous El
Capitan reef complex of West Texas formed during this interval. The Paleo-
zoic era ended with a sweeping extinction event that involved not only reef
inhabitants but also other marine organisms.
After the Paleozoic extinctions, reefs were largely absent during the
early part of the Mesozoic era. The advent of modern reefs consisting of
scleractinian corals and red and green algae occurred in the Late Triassic
period. Stromatoporoids once again occurred abundantly on reefs during
this interval; however, the role of the previously ubiquitous blue-green al-
gal stromatolites in reefs declined. Late Cretaceous reefs were often domi-
nated by conical, rudistid bivalves that developed the ability to form
frameworks and may have possessed symbiotic relationships with algae,
as do many modern corals. Rudists, however, became extinct during the
sweeping extinctions that occurred at the end of the Cretaceous period.
The reefs that were reestablished in the Cenozoic era lacked stromato-
poroids and rudists and consisted of scleractinian corals and red and green
calcareous algae. This reef type has persisted, with fluctuations, until the
present.
Study of Modern Reefs

Modern reefs are typically studied while scuba (self-contained underwater
breathing apparatus) diving, which enables observation and sampling to
a depth of approximately 50 meters. Deeper environments have been
made accessible through the availability of manned submersibles and un-
manned, remotely operated vehicles that carry mechanical samplers and
still and video cameras. The biological compositions of reef communities
are determined by census (counting) methods commonly employed by
plant ecologists. Studies of symbioses, such as that between corals and
their zooxanthellae, employ radioactive tracers to determine the transfer
of products between symbiont and host. Growth rates are measured by
staining the calcareous skeletons of living organisms with a dye, such as
Alizarin red, and then later collecting and sectioning the specimen and
568
Reefs
measuring the amount of skeleton added since the time of staining. An-
other method for determining growth is to X-ray a thin slice of skeleton
and then measure and count the yearly growth bands that are revealed on
the X-radiograph. Variations in growth banding reflect, among other fac-
tors, fluctuations in ocean temperature.
Reef sediments, which will potentially be transformed into reef lime-
stones, are examined through sieving, X-ray diffraction, and epoxy im-
pregnation and thin-sectioning. Sieving enables the determination of sedi-
ment texture, the relationships of grain sizes and abundance (which will
reflect environmental energy and the production), and erosion of grains
through biotic processes. X-ray diffraction produces a pattern that is deter-
mined by the internal crystalline structure of the sediment grains. As each
mineral possesses a unique structure, the mineralogical identity of the
sediment may be determined. Thin sections of embedded sediment or
lithified rock are examined with petrographic microscopes, which reveal
the characteristic microstructures of the individual grains. Thus, even
highly abraded fragments of coral or algae may be identified and their con-
tributions to the reef sediment determined.
Because of their typically massive nature, fossil reefs are usually stud-
ied by thin-sectioning of lithified rock samples collected either from sur-
face exposures or well cores. Reef limestones that have not undergone ex-
tensive alteration may be dated through carbon 14 dating, if relatively
young, or through uranium-series radiometric dating methods.
Reefs as Ecological Laboratories

Modern reefs serve as natural laboratories, enabling the scientists to wit-
ness and study phenomena, such as carbonate sediment production, bio-
erosion, and early cementation, that have been responsible for forming
major carbonate rock bodies in the past. The study of cores extracted from
centuries-old coral colonies shows promise for deciphering past climates
and perhaps predicting future trends. This is made possible by the fact that
the coral skeleton records variations in growth that are related to ocean
temperature fluctuations. The highly diverse modern reefs also serve as
ecological laboratories for testing models on the control of community
structure. For example, the relative importance of stability versus distur-
bance and recruitment versus predation in determining community struc-
ture is being studied within the reef setting.
Modern reefs are economically significant resources, particularly for
many developing nations in the tropics. Reefs and the associated lagoonal
sea-grass beds serve as important nurseries and habitats for many fish and
invertebrates. The standing crop of fish immediately over reefs is much
569
Reefs
higher than that of adjacent open shelf areas. Reef organisms may one
day provide an important source of pharmaceutical compounds, such as
prostaglandins, which may be extracted from gorgonians (octocorals). In
addition, research has focused upon the antifouling properties exhibited
by certain reef encrusters. Reefs also provide recreational opportunities for
snorkelers and for scuba divers, a fact that many developing countries are
utilizing to promote their tourist industries. Finally, reefs serve to protect
shorelines from wave erosion.
Because of the highly restricted environmental tolerances of reef organ-
isms, the occurrence of reefs in ancient strata enables fairly confident esti-
mation of paleolatitude, temperature, depth, salinity, and water clarity. In
addition, depth- or turbulence-related variation in growth form (mounds
in very shallow water, branches at intermediate depths, and plates at
greater depths) enables even more precise estimation of paleobathymetry
or turbulence levels. Finally, buried ancient reefs are often important reser-
voir rocks for hydrocarbons and thus are important economic resources.
W. David Liddell
See also: Defense mechanisms; Marine biomes; Ocean pollution and oil
spills; Phytoplankton.

Birkeland, Charles, ed. Life and Death of Coral Reefs. New York: Chapman
and Hall, 1997.
Cousteau, Jacques-Yves, and Philippe Diolé. Life and Death in a Coral Sea.
Translated by J. F. Bernard. Garden City, N.Y.: Doubleday, 1971.
Darwin, Charles. The Structure and Distribution of Coral Reefs. 1851. Reprint.
Berkeley: University of California Press, 1962.
Davidson, Osha Gray. The Enchanted Braid: Coming to Terms with Nature on
the Coral Reef. New York: Wiley, 1998.
Frost, S. H., M. P. Weiss, and J. B. Sanders, eds. Reefs and Related Carbonates:
Ecology and Sedimentology. Tulsa, Okla.: American Association of Petro-
leum Geologists, 1977.
Goreau, Thomas F., et al. “Corals and Coral Reefs.” Scientific American 241
(August, 1979): 16.
Jones, O. A., and R. Endean, eds. Biology and Geology of Coral Reefs. 4 vols.
New York: Academic Press, 1973-1977.
Kaplan, Eugene H. A Field Guide to Coral Reefs of the Caribbean and Florida,
Including Bermuda and the Bahamas. Boston: Houghton Mifflin, 1988.
Keating, Barbara H., et al., eds. Seamounts, Islands, and Atolls. Washington,
D.C.: American Geophysical Union, 1987.
570
Reefs
Köhler, Annemarie, and Danja Köhler. The Underwater Explorer: Secrets of a

Blue Universe. New York: Lyons Press, 1997.
National Geographic Society. Jewels of the Caribbean Sea: A National Geo-
graphic Special. Video. Washington, D.C.: Author, 1994.
Newell, Norman D. “The Evolution of Reefs.” Scientific American 226 (June,
1972): 12, 54-65.
Wood, Rachel. Reef Evolution. New York: Oxford University Press, 1999.
571
REFORESTATION
Reforestation is the growth of new trees in an area that has been cleared for human
activities. It can occur naturally or be initiated by people.
M any areas of the eastern United States, such as the New England re-
gion, reforested naturally in the nineteenth and early twentieth cen-
turies after farmland that had been abandoned was allowed to lie fallow
for decades. After an area has been logged, environmentalists, as well as
the commercial logging industry, advocate planting trees rather than wait-
ing for natural regrowth because the process of natural regeneration can be
both slow and unpredictable. In natural regeneration, the mixture of trees
in an area may differ significantly from the forest that preceded it. For ex-
ample, when nineteenth century loggers clear-cut the white pine forests of
the Great Lakes region, many logged-over tracts grew back primarily in
mixed hardwoods.
Land that has been damaged by industrial pollution or inefficient agri-
cultural practices sometimes loses the ability to reforest naturally. In some
regions of Africa, soils exposed by slash-and-burn agriculture contain high
levels of iron or aluminum oxide. Without a protective cover of vegetation,
even under cultivation, soil may undergo a process known as laterization.
Laterite is a residual product of rock decay that makes soil rock-hard. Such
abandoned farmland is likely to remain barren of plant life for many years.
In polluted areas such as former mining districts, native trees may not be
able to tolerate the toxins in the soil; in these cases, more tolerant species
must be introduced.
Safeguarding Timber Resources

Reforestation differs from tree farming in that the goal of reforestation is
not always to provide woodlands for future harvest. Although tree farm-
ing is a type of reforestation (trees are planted to replace those that have
been removed), generally only one species of tree is planted, with explicit
plans for its future harvest. The trees are seen first as a crop and only inci-
dentally as wildlife habitat or a means of erosion control.
As foresters have become knowledgeable about the complex interac-
tions within forest ecosystems, however, tree farming methods have begun
to change. Rather than monocropping (planting only one variety of tree),
the commercial forest industry has begun planting mixed stands. Trees
572
Reforestation
that possessed no commercial value, once considered undesirable weed

trees, are now recognized as nitrogen fixers necessary for the healthy
growth of other species. In addition to providing woodlands for possible
use in commercial forestry, goals of reforestation include wildlife habitat
restoration and the reversal of environmental degradation.
Early Efforts
Reforestation to replace trees removed for commercial purposes has been
practiced in Western Europe since the late Middle Ages. English monarchs,
including Queen Elizabeth I, realized that forests were a vanishing re-
source and established plantations of oaks and other hardwoods to ensure
a supply of ship timbers. Similarly, Sweden created a corps of royal forest-
ers to plant trees and watch over existing woodlands. These early efforts at
reforestation were inspired by the reduction of a valuable natural resource.
By the mid-nineteenth century it was widely understood that the removal
of forest cover contributes to soil erosion, water pollution, and the disap-
pearance of many species of wildlife.
Clear-cutting removes all the trees on a tract of land, leaving none standing. At
one time a standard practice in logging, it has become one of the most controversial
harvesting techniques used in modern logging. With its windrows of slash and de-
bris, a clear-cut tract of land may appear as though a catastrophic event has devas-
tated the landscape. (PhotoDisc)
573
Reforestation
The laborious process of

planting new trees, combined
with the time required for those
trees to grow, highlights the
need to eliminate clear-cutting
as a means of harvesting
timber. Reforestation also
requires careful reconstruction
of the forest community in all
its variety, in order to provide
habitat for many forms of
wildlife. Simple “tree
farming,” by contrast, is aimed
at growing stands of a single
species of tree for later
harvesting. (PhotoDisc)
Ecological and Environmental Aspects

Water falling on hillsides made barren by clear-cutting timber washes
away topsoil and causes rivers to choke with sediment, killing aquatic life.
Without trees to slow the flow of water, rain can also run off slopes too
quickly, causing rivers to flood. For many years, soil conservationists ad-
vocated reforestation as a way to counteract the ecological damage caused
by erosion.
In the mid-twentieth century, scientists established the vital role that
trees, particularly those in tropical rain forests, play in removing carbon di-
oxide from the earth’s atmosphere through the process of photosynthesis.
Carbon dioxide is a greenhouse gas: It helps trap heat in the atmosphere.
As forests disappear, the risk of global warming—caused in part by an in-
crease in the amount of carbon dioxide in the atmosphere—becomes
greater. Since the 1980’s, scientists and environmental activists concerned
about global warming have joined foresters and soil conservationists in
urging that for every tree removed anywhere, whether to clear land for de-
velopment or to harvest timber, replacement trees be planted. As the area
covered by tropical rain forests shrinks in size, the threat of irreversible
damage to the global environment becomes greater.
574
Reforestation
Reforestation Programs
In 1988 American Forests, an industry group, established the Global ReLeaf
program to encourage reforestation efforts in an attempt to combat global
warming. In addition to supporting reforestation efforts by government
agencies, corporations, and environmental organizations, Global ReLeaf
and similar programs encourage people to practice reforestation in their
own neighborhoods. Trees serve as a natural climate control, helping to
moderate extremes in temperature and wind. Trees in a well-landscaped
yard can reduce a homeowner’s energy costs by providing shade in the
summer and serving as a windbreak during the winter. Global ReLeaf is
one of many programs that support reforestation efforts.
Arbor Day, an annual day devoted to planting trees for the beautifica-
tion of towns or the forestation of empty tracts of land, was established in
the United States in 1872. The holiday originated in Nebraska, a prairie
state that seemed unnaturally barren to homesteaders used to eastern
woodlands. Initially emphasizing planting trees where none had existed
before, Arbor Day is observed in U.S. public schools to educate young peo-
ple about the importance of forest preservation. Organizations such as the
National Arbor Day Foundation provide saplings (young trees) to schools
and other organizations for planting in their own neighborhoods.
See also: Biodiversity; Biomass related to energy; Conservation biology;

Deforestation; Endangered plant species; Erosion and erosion control; For-
est fires; Forest management; Forests; Grazing and overgrazing; Integrated
pest management; Multiple-use approach; Old-growth forests; Rain for-
ests; Restoration ecology; Species loss; Sustainable development; Urban
and suburban wildlife; Wildlife management.

Cherrington, Mark. Degradation of the Land. New York: Chelsea House,
1992.
TreePeople, with Andy Lipkis and Katie Lipkis. The Simple Act of Planting a
Tree: A Citizen Forester’s Guide to Healing Your Neighborhood, Your City, and
Your World. Los Angeles: St. Martin’s Press, 1990.
Weiner, Michael. Plant a Tree: Choosing, Planting, and Maintaining This Pre-
cious Resource. New York: Wiley, 1992.
575
REPRODUCTIVE STRATEGIES
Types of ecology: Behavioral ecology; Population ecology
Reproductive strategies are a set of attributes involved in an organism’s maximiz-

ing its reproductive success. Theoretical and experimental studies of reproductive
strategies reveal why various reproductive patterns have evolved.
T he concept of reproductive strategies is closely related to that of natu-

ral selection. Natural selection results in the individuals within a pop-
ulation, under a given set of environmental circumstances, being more likely
to pass on their genes to future generations. By this process, the gene pool
(genetic makeup) of the population is altered over time. An organism’s fit-
ness can be assessed by evaluating two key characteristics: survival and re-
productive success. The organism’s reproductive strategy, then, is the
blend of traits enabling it to have the highest overall reproductive success.
Application of the term “reproductive strategy” has also been extended to
describe patterns beyond individual organisms: populations, species, and
even entire groups of similar species, such as carnivorous mammals.
Examination of reproductive strategies is part of the larger study of life-
history evolution, which attempts to understand why a given set of basic
traits has evolved. These traits include not only those pertaining to repro-
duction but also those such as body size and longevity. To consider a repro-
ductive strategy appropriately, one must view it within the context of the
organism’s overall life history, precisely because these traits (particularly
body size) often affect reproductive traits. One should also evaluate the
role that the organism’s ancestry plays in these processes. A species’ evolu-
tionary history can have a profound effect on its current attributes.
Reproductive Traits and Behaviors

A reproductive strategy consists of a collection of basic reproductive traits,
including litter, or “clutch,” size (the number of offspring produced per
birth), the number of litters per year, the number of litters in a lifetime, and
the time between litters, gestation, or pregnancy length. The age of the
mother’s first pregnancy is also a consideration. Another trait is the degree
of development of the young at birth. In different species, mothers put
varying levels of time and energy into the production of either relatively
immature, or altricial, offspring or offspring that are well developed, or
precocial.
Reproductive strategies also consist of behavioral elements, such as the
576
Reproductive strategies
mating system and the amount of parental care. Mating systems include
monogamy (in which one male is mated to one female) and polygamy (in
which an individual of one sex is mated to more than one from the other).
The type of polygamy when one male mates with several females is called
polygyny; the reverse is known as polyandry.
Finally, physiological events such as those involved in ovulation (what
happens when the egg or eggs are shed from the ovary) may also be used
to characterize a reproductive strategy. Some mammals are spontane-
ous ovulators. Females shed their eggs during the reproductive cycle
without any physical stimulation. Other mammalian species are induced
ovulators—a female ovulates only after being physically stimulated by a
male during copulation. These patterns, induced and spontaneous ovula-
tion, may be regarded as alternate reproductive strategies, each enabling a
type of species to reproduce successfully under certain conditions.
The overall effectiveness of a reproductive strategy is important to con-
sider with respect to the relative success of the offspring (even those in fu-
ture generations) in leaving their own descendants. A sound reproductive
strategy results in increased fitness. An organism’s fitness as it affects the
population’s gene pool may not be adequately assessed until several gen-
erations have passed.
The r and K Selection Model

The model of r and K selection is the most widely cited description of how
certain reproductive traits are most effective under certain environmental
conditions. To appreciate this model, an understanding of elementary pop-
ulation dynamics is needed. At the early stages of a population’s growth,
the rate of addition of new individuals (designated r) tends to be slow. Af-
ter a sufficient number of individuals is reached, the growth rate can in-
crease sharply, resulting in a boom phase. In most environments, however,
unrestrained growth cannot continue indefinitely. Critical resources—
food, water, and protective cover—become more scarce as the environ-
ment’s carrying capacity (K) is approached. Carrying capacity is the maxi-
mal population size an area can support. When the population approaches
this level, growth rate slows as individuals now have fewer resources to
convert into the production of new offspring.
This pattern is defined as density-dependent population growth—the
density or number of individuals per area that influences its growth. This
description of population dynamics is also referred to as logistic growth
and was conceived by the Belgian mathematician Pierre-François Verhulst
in the early nineteenth century. It has successfully described population
growth in many species.
577
The r and K selection model was presented by Robert H. MacArthur

and Edward O. Wilson in their influential book, The Theory of Island Bioge-
ography (1967). They argued that in the early phase of a population’s
growth, individuals should evolve traits associated with high reproduc-
tive output. This enables them to take advantage of the relatively plentiful
supply of food. The evolution of such traits is called r selection, after the
high population growth rates occurring during this phase. They also sug-
gested that, as the carrying capacity was approached, individuals would
be selected that could adjust their lives to the now reduced circumstances.
This process is called K selection. Such individuals should be more effi-
cient in the conversion of food into offspring, producing fewer young than
those living in the population’s early phase. In a sense, a shift from produc-
tive to efficient individuals occurs as the population grows.
Other biologists, most notably Eric Pianka, have extended this concept
of r and K selection to entire species rather than only to individuals at dif-
ferent stages of a population’s growth. Highly variable or unpredictable
climates commonly create situations in which population size is first di-
minished but then grows rapidly. Species commonly occurring in such en-
vironments are referred to as r strategists. Those living in more constant,
Male seals can successfully defend areas containing from eighty to a hundred fe-
males from other males. Very dense clusters of females, however, attract too many
males for one male to monopolize. When this happens, the largest male typically
dominates the rest and maintains disproportionate access to females. (PhotoDisc)
578
relatively predictable climates are less likely to go through such an explo-

sive growth phase. These species are considered to be K strategists. Ac-
cording to this scheme, an r strategist is characterized by small body size,
rapid development, high rate of population increase, early age of first re-
production, a single or few reproductive events, and many small offspring.
The K strategist has the opposite qualities—large size, slower develop-
ment, delayed age of first reproduction, repeated reproduction, and fewer,
larger offspring.
Various combinations of r and K traits may occur in a species, and few
are entirely r- or K-selected. Populations of the same species commonly oc-
cupy different habitats during their lives or across their geographic ranges.
An organism might thus shift strategies in response to environmental
changes—they may, however, be constrained by their phylogeny or ances-
try in the degree to which their strategies are flexible.
Criticism of the Model

Because the r and K model of reproductive strategies seems to explain pat-
terns observed in nature, it has become widely accepted. It has also met
with considerable criticism. Charges against it include arguments that the
logistic population-growth model (on which the r and K strategies model
is based) is too simplistic. Another is that cases of r and K selection have
not been adequately tested. Ecologist Mark Boyce has persuasively argued
that for the r and K model to be most useful it must be viewed as a model of
how population density affects life-history traits. Within this framework,
also called density-dependent natural selection, the concept of r and K
selection remains true to the one that MacArthur and Wilson originally
proposed. Boyce suggests that the ability of r and K selection to explain re-
productive strategies will have the best chance of being realized when ap-
proached in this fashion.
In addition to the r and K model, there are many other ways of describ-
ing reproductive strategies. For example, some species, such as the Chi-
nook salmon, are semelparous: They reproduce only once before dying.
The alternate is to be iteroparous—in which an organism experiences two
or more reproductive events over its life span. If juvenile death rates
are high, an individual might be better off reproducing on several occa-
sions rather than only once. (This reproductive strategy is referred to as
“bet-hedging.”) Finally, it has also been useful to evaluate reproductive
strategies based on the proportion of energy that goes into reproduction
relative to that devoted to all other body functions. This mode of analy-
sis addresses such considerations as reproductive effort and resource allo-
cation.
579
Studying Reproductive Strategies

Initially, one who studies the reproductive strategy of an organism should
attempt to characterize its reproduction fully. The sample examined must
be representative of the population under consideration—it should ac-
count for the variability of the traits being measured. Studies can involve
any of several approaches. Short-term laboratory studies can uncover
some hard-to-observe features, but there is no substitute for long-term
field research. By studying an organism’s reproduction in nature, a biolo-
gist has the best chance of determining how its reproduction is shaped by
an environment. If the research is performed over several seasons or years,
patterns of variability can be better understood. This is important in deter-
mining how the physical environment influences reproductive traits.
After data have been systematically collected, it might then be possible
to characterize a reproductive strategy. Imagine that a mouse population
becomes established in a previously uninhabited area and that the popula-
tion has a high reproductive rate (it produces large litters). The young de-
velop quickly and produce many young themselves. Because of this com-
bination of circumstances, one might consider the reproductive strategy to
be r-selected since the population has a high reproductive output in an un-
exploited area. Though the concept of r and K strategies is problematic, it
still is common to typify a strategy as r- or K-selected based upon this ap-
proach.
Because a reproductive strategy needs to be seen as part of an organ-
ism’s overall life history, however, other things should be measured to un-
derstand it fully. These may include the life span and population attributes
such as survival patterns. Values should be taken for different age groups
to characterize the population’s strategy. Correlational analysis is a statisti-
cal procedure that is used to evaluate reproductive strategies. Through
such a methodology, one assesses the degree of association between two
variables or factors. This may involve relationships between two repro-
ductive variables or between a reproductive and an environmental vari-
able—for example, to determine if there is a significant correlation be-
tween litter size and decreasing body size in mammals. If one were found
to occur, the conclusion that smaller species typically have larger litters
might be drawn, which is, in fact, true. Such an analysis enables the charac-
terization of a change in reproductive strategy based on body size. Simply
establishing a correlation does not prove that causation has occurred—it
does not automatically mean that one factor is responsible for the expres-
sion of the other.
Multivariate statistical procedures are also used to analyze reproduc-
tive strategies. These allow the determination of how groups of reproduc-
580
tive traits are associated and of how they can be explained by several fac-
tors. One might determine that a certain bird species produces its greatest
number of young, and that the young grow most rapidly, at northern loca-
tions having high snow levels. Such an approach is often needed in dealing
with reproductive strategies—a combination of traits typically requires ex-
planation.
Reproduction and Survivial

The characterization of an organism’s reproductive strategy involves more
than an understanding of reproductive traits. There is a successful process
by which offspring are produced, and reproductive success is one of the
two principal measures of fitness—the other is survival. Because a success-
ful reproductive strategy ultimately results in high fitness, any discussion
of these strategies bears directly on issues of natural selection and evolu-
tion.
An organism’s reproductive strategy represents perhaps the most sig-
nificant way the organism is adapted to its environment. A successful re-
productive strategy represents a successful mode of passing genes on to
the next generation, so traits associated with a reproductive strategy are
under intense natural selection pressure. If environmental conditions
change, the original strategy may no longer be as successful. To the extent
that an organism can shift its reproductive strategy as circumstances
change, its genes will persist.
The study of reproductive strategies has helped scientists understand
why certain modes of reproduction occur, based upon observations of a
species itself and of its environment. An understanding of reproductive
strategies may also be of some practical use. An organism’s reproduction
directly influences its population dynamics.
If an animal has small litters and is at an early age at first reproduction,
its population should grow at a concomitantly high rate. These and other
components of reproduction may strongly affect a species’ population
growth. A knowledge of how reproduction influences population dynam-
ics can be important in wildlife management activities, which can range
from strict preservation efforts to overseeing trophy hunting.
Samuel I. Zeveloff
See also: Adaptive radiation; Biodiversity; Biogeography; Clines, hybrid

zones, and introgression; Convergence and divergence; Demographics;
Displays; Ethology; Extinctions and evolutionary explosions; Gene flow;
Genetic diversity; Genetic drift; Human population growth; Insect societ-
ies; Natural selection; Nonrandom mating, genetic drift, and mutation;
581
Population analysis; Population fluctuations; Population genetics; Popula-

tion growth; Punctuated equilibrium vs. gradualism; Speciation; Terri-
toriality and aggression.

Austin, C. R., and R. V. Short, eds. The Evolution of Reproduction. New York:
Boyce, Mark S. “Restitution of r- and K-Selection as a Model of Density-
Dependent Natural Selection.” Annual Review of Ecology and Systematics
15 (1984): 427-447.
Clutton-Brock, T. H., F. E. Guiness, and S. D. Albon. Red Deer: Behavior and
Ecology of Two Sexes. Chicago: University of Chicago Press, 1982.
Ferraris, Joan D., and Stephen R. Palumbi, eds. Molecular Zoology: Ad-
vances, Strategies, and Protocols. New York: Wiley-Liss, 1996.
MacArthur, Robert H., and Edward O. Wilson. The Theory of Island Biogeog-
raphy. Princeton, N.J.: Princeton University Press, 1967.
Pianka, Eric R. Evolutionary Ecology. 6th ed. New York: Harper & Row,
2000.
Wrangham, Richard W., W. C. McGrew, Frans B. M. De Waal, and Paul G.
Heltne, eds. Chimpanzee Cultures. Cambridge, Mass.: Harvard Univer-
sity Press, 1996.
582
RESTORATION ECOLOGY
Restoration ecology is concerned with converting ecosystems that have been modi-
fied or degraded by human activity to a state approximating their original condi-
tion.
F ederal laws often dictate ecological restoration following strip mining,

construction, and other activities that alter the landscape. As a part of
the management of natural areas that have been disturbed to some degree,
several options are available. One is to do nothing but protect the property,
allowing nature to take its course. In the absence of further disturbances,
one would expect the area to undergo the process of ecological succession.
Theoretically, an ecosystem similar to that typical of the region, and includ-
ing an array of organisms, would be expected to return.
One might, therefore, ask why ecological restoration is mandated. For
one thing, succession is often a process requiring long periods of time. As
an example, the return of a forest following the destruction of the trees and
the removal of the soil would require more than one century. Also, ecosys-
tems resulting from succession may be lacking in species typical of the re-
gion. This is true when succession is initiated in an area where many exotic
(alien) species are present or where certain native species have been elimi-
nated. Succession can produce a new ecosystem with a biodiversity com-
parable to the original one only if there is a local source of colonizing ani-
mals and seeds of native plants. Also, satisfactory recovery by succession
is unlikely if the soil has been heavily polluted by heavy metals or other
substances caused by industrial land use.
The Restoration Process

Once it has been decided that a given ecosystem is to be restored, success
requires that a plan be designed and followed. Although the specifics may
vary greatly, all restoration projects should follow five basic steps: Envi-
sion the end result, consult relevant literature and solicit the advice of spe-
cialists, remove or mitigate any current disturbances to the site, rehabili-
tate the physical habitat, and restore indigenous plants and animals.
Much can be learned from restoration projects that have been con-
ducted in various parts of the world involving a wide variety of ecosys-
tems. A classic ecological restoration of a prairie was conducted in Wiscon-
sin beginning in the 1930’s. Because most North American prairies have
583
Restoration ecology
Restoration ecology enlists human intervention to return habitats and ecosystems

to their former state, particularly when natural processes such as ecological suc-
cession would take far longer than the original destruction of the community. Such
destruction is often the result of human activity such as clear-cutting, strip-min-
ing, large-scale agriculture, or other development. (PhotoDisc)
been converted to agricultural uses, many opportunities exist for prairie

restoration. In such projects it is often necessary to eliminate exotic plants
by mechanical means or by application of herbicides. Native prairie
grasses and forbs can be established by transplantation or from seed. It
may also be necessary to introduce native fauna from nearby areas. Peri-
odic prescribed burning is often necessary to simulate natural fires com-
mon in prairies.
After decades of loss of wetlands in the United States, governmental
policy is now “no net loss.” Thus, when a wetland is destroyed by develop-
ment, it is required that a new wetland be created as compensation. Before
introducing native biota, it is necessary to alter the hydrology of the new
site.
Thomas E. Hemmerly
See also: Biological invasions; Conservation biology; Deforestation; En-

dangered animal species; Endangered plant species; Erosion and erosion
control; Forest management; Grazing and overgrazing; Integrated pest
management; Invasive plants; Landscape ecology; Multiple-use approach;
584
Restoration ecology
Old-growth forests; Reforestation; Succession; Sustainable development;

Urban and suburban wildlife; Waste management; Wildlife management;
Zoos.

Harper, David. Eutrophication of Freshwaters: Principles, Problems, and Resto-
ration. London: Chapman & Hall, 1992.
Hey, Donald L., and Nancy S. Philippi. A Case for Wetland Restoration. New
York: Wiley, 1999.
585
SAVANNAS AND DECIDUOUS
TROPICAL FORESTS
Savannas are areas of continuous grass or sedge cover beneath trees that range
from scattered, twisted, and gnarled individuals to open woodlands. Deciduous
tropical forests have continuous to open forest cover and undergo a leafless period
during a seasonally lengthy dry season.
W here the annual rainfall in tropical regions is less than 2,000 millime-
ters and three to six months out of the year are dry, savannas and de-
ciduous forests are common. Deciduous tropical forests often occur where
the annual rainfall is less than that of savannas. Together, the two biomes
are referred to here as the dry tropical biome.
A pronounced pattern of seasonally wet and dry periods is the most im-
portant factor affecting the distribution of these types of plant cover.
Higher soil fertility favors forest over grasses and savanna such as in the
cerrado of Brazil, which occurs only on certain geological formations and
low-nutrient soils. Fire has been a dominant feature of these biomes, and
human influences—fires, agriculture, and grazing of animals—have inter-
acted with climate to produce a varied landscape.
The dry tropical biome is most geographically widespread on the conti-
nents of Africa, South America, and Australia, with smaller enclaves in
Asia. The world’s largest expanses of dry forest—the Brachystegia wood-
land across Central Africa, the cerrado (savanna) and caatinga (dry forest)
of the Amazon basin, and much of interior Australia—are notable exam-
ples. “Elephant grass savanna,” with tall grasses up to 4 meters tall and
scattered trees, occurs exclusively in Africa. In the West Indies, dry forest
occurs in rain-shadow zones on the leeward sides of islands affected by the
tradewinds.
Plant Adaptations and Diversity of Life-Forms

As the rainfall decreases below 2,000 millimeters, and especially below
1,000 millimeters, the height of the forest decreases and the proportion of
trees that are deciduous increases. In the dry tropics, leaf fall occurs in
response to drought, and therefore the lengthy dry season becomes a selec-
tive pressure to which plants have adapted. Tree leaves tend to be com-
pound, with small leaflets that help plants exchange heat with their sur-
roundings better than large, simple leaves; rates of leaf respiration and
586
Savannas and deciduous tropical forests
transpiration are thereby reduced. Sclerophyllous leaves are common, aid-

ing in moisture retention, and the drier, more open woodlands may have
cacti or other succulents.
The dry forest is far less species-rich than the rain forest, but the di-
versity of life-forms and the proportion of endemics are greater. For exam-
ple, dry forests may contain xerophytic (dry-adapted) evergreens, either
obligatively or facultatively deciduous trees, trees with photosynthetic
bark, plants that use the Crassulacean acid metabolism (CAM) photosyn-
thesis as well as C3 and C4 dicots, grasses, bromeliads, lianas, epiphytes,
and hemiparasites. Trees from Fabaceae (the legume family) are the most
well-represented family among trees.
Dry forests contain a higher proportion of wind-dispersed species than
wetter forests, and many trees will have their flowering and fruiting
controlled by the duration and intensity of the dry season. Synchronous
flowering within and among species is common, and many produce seed
during the dry season. Flowers are often conspicuous and visited by spe-
cialized pollinators such as hawkmoths, bats, and bees.
It is incorrect to generalize about savannas and dry tropical forests be-
cause, although they both occur in the drier tropics, the two vegetation
Savannas are landscapes of dense grass and scattered trees, such as these yellow fe-
ver trees growing in Nakura National Park in Kenya. Common on the continent of
Africa, savannas are also found in India, Australia, and the northern part of South
America. (Corbis)
587
types occur in different habitats and are adapted differently to their respec-
tive environments.
Trees of the cerrado in northeast Brazil are deeply rooted, tap ground-
water, and have high rates of transpiration. Drought here is atmospheric,
as water is always available below two meters of soil depth. The deciduous
caatinga of central Brazil, however, receives only 500 millimeters of rain
yearly, and transpiration of trees is low. Here, trees suffer significant water
deficits during the long, dry season, are truly xerophytic, and exhibit clas-
sic adaptations to drought.
Trees of the cerrado have a number of adaptations that confer resistance
to fire. These include a thick, corky bark, the ability to form adventitious
roots from buds on roots following the burning of the stem, and the
cryptophyte or hemicryptophyte life-form (cryptophytes produce buds
underground). Many herbaceous species are induced to flower by fire.
Human Impacts and Conservation

Fires have occurred in the Brazilian cerrado for thousands of years, based
on carbon 14 dating of charcoal fragments. Fire is thus an environmen-
tal factor to which the vegetation has become adapted. Yet, the human
influence has been to increase the incidence of fire. The cerrado has
changed as a result to a more open form of plant cover with fewer trees
and shrubs. In addition, timber extraction, charcoal production, and ranch-
ing have altered the savanna landscape. The ability of belowground or-
gans to survive such types of disturbance has increased the ability of the
cerrado to persist. Yet it is estimated that 50 percent of the cerrado has been
destroyed, much of this the result of clearing for agriculture since the
1960’s.
Because of better soils and fewer pests, humans in tropical areas of Cen-
tral America have mostly chosen the dry and moist life zones as places to
live rather than the wetter rain-forest zones. As a result, dry forest ecosys-
tems have been subject to massive disturbance. Today, only a small fraction
of the original dry forest remains. Fire has been used as a means of clear-
ing the forest for farming, but, unlike the savanna, the dry forest is not
adapted to fire. At Guanacaste, Costa Rica, a well-known tropical conser-
vation project, restoration of dry forest is dependent on controlling annual
fires set by farmers and ranchers and supporting the return of forest vege-
tation to dry areas. In Africa, large areas of dry forest are burned annually
by farmers, and areas of dense, dry forest have been converted to more
open forest or even savanna. Sustainable land-use systems are urgently
needed for dry tropical regions.
Allan P. Drew
588
See also: Biomes: determinants; Biomes: types; Forests; Grasslands and

prairies; Habitats and biomes; Rangeland; Reforestation.

Allen, W. Green Phoenix: Restoring the Tropical Forests of Guanacaste, Costa
Rica. New York: Oxford University Press, 2001.
Bullock, S. H., H. A. Mooney, and E. Medina. Seasonally Dry Tropical Forests.
Rizzini, C. T., A. F. C. Filho, and A. Houaiss. Brazilian Ecosystems. Rio de Ja-
neiro: ENGE-RIO, Index Editora, 1988.
Silver, Donald M. African Savanna. New York: McGraw-Hill, 1997.
589
SLASH-AND-BURN AGRICULTURE
Slash-and-burn agriculture, also called swidden agriculture, is a practice in which

forestland is cleared and burned for use in crop and livestock production. While
yields are high during the first few years, they rapidly decline in subsequent years,
leading to further clearing of nearby forestland.
S lash-and-burn agriculture has been practiced for many centuries among

people living in tropical rain forests. Initially, this farming system in-
volved small populations. Therefore, land could be allowed to lie fallow
(unplanted) for many years, leading to the full regeneration of the second-
ary forests and hence a restoration of the ecosystems. During the second
half of the twentieth century, however, several factors led to drastically re-
duced fallow periods. In some places such fallow systems are no longer in
existence, resulting in the transformation of forests into shrub and grass-
lands, negative effects on agricultural productivity for small farmers, and
disastrous consequences to the environment.
Among the factors that have been responsible for reduced or nonexistent
fallow periods are increased population in the tropics, increased demand for
wood-based energy, and, perhaps most important, the increased worldwide
demand for tropical commodities during the 1980’s and 1990’s, especially
for products such as palm oil and natural rubber. These last two factors
have helped industrialize slash-and-burn agriculture, which was practiced
for centuries mainly by small farmers. Ordinarily, small farmers are able to
control their fires so that they are similar to a small forest fire triggered by
lightning in the northwestern or southeastern United States. However, the
continued reduction in fallow periods, coupled with increased burning by
subsistence farmers and large agribusiness, especially in Asia and Latin
America, is resulting in increased environmental concern.
While slash-and-burn agriculture seldom takes place in temperate re-
gions, some agricultural burning occurs in the Pacific Northwest of the
United States, where it is estimated that three thousand to five thousand
agricultural fires are set each year in Washington State alone. These fires
also create problems for human health and the environment.
Habitat Fragmentation
One of the most easily recognized results of slash-and-burn agriculture is
habitat fragmentation, which leads to a significant loss of the vegetation
590
Slash-and-burn agriculture
needed for the maintenance of effective gaseous exchange in tropical re-

gions and throughout the world. For every acre of land lost to slash-and-
burn agriculture, 10 to 15 acres (4 to 6 hectares) of land are fragmented, re-
sulting in the loss of habitat for wildlife, plant species, and innumerable
macro- and microorganisms yet to be identified. This also creates problems
for management and wildlife conservation efforts in parts of the world
with little or no resources to feed their large populations. Fragmentation
has also led to intensive discussions on global warming. While slash-and-
burn agriculture by itself is not completely responsible for global warm-
ing, the industrialization of the process could make it a significant compo-
nent of the problem, as more and more vegetation is fragmented.
Human Health
The impact of slash-and-burn agriculture on human health and the envi-
ronment is best exemplified by the 1997 Asian fires that resulted from such
practices. Monsoon rains normally extinguish the fires set by farmers, but
a strong El Niño weather phenomenon delayed the expected rains, and the
fires burned out of control for months. Thick smoke caused severe health
problems. It is estimated that more than 20 million people in Indonesia
alone were treated for asthma, bronchitis, emphysema, and eye, skin, and
cardiovascular problems as a result of the fires. Similar problems have
been reported for smaller agricultural fires.
Three major problems are associated with air pollution: particulate mat-
ter, pollutant gases, and volatile organic compounds. Particulate com-
pounds of 10 microns or smaller that are inhaled become attached to the al-
veoli and other blood cells, resulting in severe illness. Studies by the U.S.
Environmental Protection Agency (EPA) and the University of Washing-
ton indicate that death rates associated with respiratory illnesses increase
when fine particulate air pollution increases. Meanwhile, pollutant gases
such as carbon monoxide, nitric oxide, nitrogen dioxide, and sulfur diox-
ide become respiratory irritants when they combine with vapor to form
acid rain or fog. Until the Asian fires, air pollutants stemming from the
small fires of slash-and-burn agriculture that occur every planting season
often went unnoticed. Thus, millions of people in the tropics experience
environmental health problems because of slash-and-burn agriculture that
are never reported.
Soil and Water Quality

The loss of vegetation that follows slash-and-burn agriculture causes an in-
creased level of soil erosion. The soils of the humid tropics create a hard
pan underneath a thick layer of organic matter. Therefore, upon the re-
591
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moval of vegetation cover, huge areas of land become exposed to the tor-
rential rainfalls that occur in these regions. The result is severe soil erosion.
As evidenced by the impact of Hurricane Mitch on Honduras during 1998,
these exposed lands can give rise to large mudslides that can lead to signif-
icant loss of life. While slash-and-burn agriculture may not be the ultimate
cause for sudden mud slides, it does predispose these lands to erosional
problems.
Associated with erosion is the impact of slash-and-burn agriculture
on water quality. As erosion continues, sedimentation of streams in-
creases. This sedimentation affects stream flow and freshwater discharge
for catchment-area populations. Mixed with the sediment are minerals
such as phosphorus and nitrogen-related compounds that enhance algal
growth in streams and estuaries, which depletes the supply of oxygen that
aquatic organisms require to survive. Although fertility is initially in-
creased on noneroded soils, nutrient deposition and migration into drink-
ing water supplies continues to increase.
Controlling Slash-and-Burn Agriculture

Given the fact that slash-and-burn agriculture has significant effects on the
environment not only in regions where it is the mainstay of the agricultural
592
systems but also in other regions of the world, it has become necessary to
explore different approaches to controlling this form of agriculture. How-
ever, slash-and-burn agriculture has evolved into a sociocultural liveli-
hood; therefore, recommendations must be consistent with the way of life
of a people who have minimal resources for extensive agricultural sys-
tems.
Among the alternatives are new agroecosystems such as agroforestry
systems and sustainable agricultural systems that do not rely so much on
the slashing and burning of forestlands. These systems allow for the culti-
vation of agronomic crops and livestock within forest ecosystems. This
protects soils from being eroded. Another possibility is the education of
small rural farmers, absentee landlords, and big agribusiness concerns in
developing countries to understand the environmental impact of slash-
and-burn agriculture. While small rural farmers may not have the re-
sources for renovating utilized forestlands, big business can organize eco-
systems restoration, as has been done in many developed nations of the
world.
Oghenekome U. Onokpise
See also: Biopesticides; Deforestation; Erosion and erosion control; Forest

management; Forests; Global warming; Grazing and overgrazing; Multiple-
use approach; Pesticides; Rangeland; Savannas and deciduous tropical for-
ests; Sustainable development.

Jordan, C. F. An Amazonian Rain Forest: The Structure and Function of a Nutri-
ent Stressed Ecosystem and the Impact of Slash and Burn Agriculture. Boca
Raton, Fla.: CRC Press, 1990.
Simons, L. M., and M. Yamashita. “Indonesia’s Plague of Fire.” National
Geographic 194 (August, 1998): 100-120.
Terborgh, J. Diversity and the Tropical Rain Forest. New York: Scientific
American Library, 1992.
Youth, Howard. “Green Awakening in a Poor Country.” World Watch 11,
no. 5 (September/October, 1998): 28-37.
593
SOIL
Types of ecology: Agricultural ecology; Ecosystem ecology; Soil ecology
Soils are complex chemical factories. Regardless of the type of soil—and twelve
types of soil are identified by the U.S. Department of Agriculture—chemical pro-
cesses such as plant growth, organic decay, mineral weathering, and water purifi-
cation, as well as living organisms, constitute the ecosystem commonly referred to
as soil.
S oil chemistry has been studied as long as there has been sustainable ag-
riculture. Although they did not recognize it as such, those first suc-
cessful farmers who plowed under plant stalks, cover crops, or animal
wastes were actively managing the soil chemistry of their fields. These
early farmers knew that to have productive farms in one location season
after season, they had to return something to the soil.
It is now understood that soil chemistry is a complex of chemical and
biochemical reactions. The most obvious result of this complex of reactions
is that some soils are very fertile whereas other soils are not. Soil itself is a
unique environment because all the “spheres”—the atmosphere, hydro-
sphere, geosphere, and biosphere—are intimately mixed there. For this
reason, soil and soil chemistry are extremely important.
Rock Weathering
Soil chemistry begins with rock weathering. The minerals composing a
rock exposed at the earth’s surface are continually bathed in a shower of
acid rain—not necessarily polluted rainwater but naturally occurring acid
rain. Each rain droplet forming in the atmosphere absorbs a small amount
of carbon dioxide gas. Some of the dissolved carbon dioxide reacts with the
water to form a dilute solution of carbonic acid. A more concentrated solu-
tion of carbonic acid is found in any bottle of sparkling water.
Most of the common rock-forming minerals, such as feldspar, will react
slowly with rainwater. Some of the chemical elements of the mineral, such
as sodium, potassium, calcium, and magnesium, are very soluble in rain-
water and are carried away with the water as it moves over the rock sur-
face. Other chemical elements of the mineral, such as aluminum, silicon,
and iron, are much less soluble. Some of these elements are dissolved in the
water and carried away; most, however, remain near the original weather-
ing, where they recombine into new, more resistant minerals. Many of the
new minerals are of a type called clays.
594
Soil
Clay minerals tend to be very small crystals composed of layers of alu-

minum and silicon. Between the layers of aluminum and silicon atoms are
positively charged ions (cations) of sodium, potassium, calcium, and mag-
nesium. The cations hold the layers of some clays together by electrostatic
attraction. In most cases, the interlayer cations are not held very tightly.
They can migrate out of the clay and into the water surrounding the clay
mineral, to be replaced by another cation from the soil solution. This phe-
nomenon is called cation exchange.
The weathering reactions between rainwater and rock minerals pro-
duce a thin mantle of clay mineral soil. The depth to fresh, unweathered
rock is not great at first, but rainwater continues to fall, percolating through
the thin soil and reacting with fresh rock minerals. In this way, the weather-
Soil Horizons
O = organic debris
A = topsoil (minerals)
B = subsoil
(clay, iron oxide,
carbonate calcium)
C = regolith (soil base)
Bedrock
595
Soil
ing front (the line between weathered minerals and fresh rock) penetrates
farther into the rock, and the overlying soil gets thicker.
Biological Processes
Throughout the weathering process, biological processes contribute to the
pace of soil formation. In the very early stages, lichens and fungi are at-
tached to what appear to be bare rock surfaces. In reality, they are using
their own acids to “digest” the rock minerals. They absorb the elements of
the mineral they need, and the remainder is left to form soil minerals. As
the soil gets thicker, larger plants and animals begin to colonize. Large
plants send roots down into the soil looking for water and nutrients. Some
of the necessary nutrients, such as potassium, are available as exchange-
able cations on soil clays or in the form of deeper, unweathered minerals.
In either case, the plant obtains the nutrients by using its own weathering
reaction carried on through its roots. The nutrient elements are removed
from minerals and become part of the growing plant’s tissue.
Without a way to replenish the nutrients in the soil, the uptake of nutri-
ents by plants will eventually deplete the fertility of the soil. Nutrients are
returned to the soil through the death and decay of plants. Microorganisms
in the soil, such as bacteria and fungi, speed up the decay. Since the bulk of
the decaying plant material is found at the surface (the dead plant’s roots
also decay), most of the nutrients are released to the surface layer of the
soil. Some of the nutrients are carried down to roots deep in the soil by in-
filtrating rainwater. Most of the nutrients, however, are removed from the
water by the shallow root systems of smaller plants. The deeper roots of
typically large plants can mine the untapped nutrients at the deep, rela-
tively unweathered soil-rock boundary.
The soil and its soil chemistry are now well established, with plants
growing on the surface and their roots reaching toward mineral nutrients
at depth. Water is flowing through the soil, carrying dissolved nutrients
and the soluble by-products of weathering reactions.
Soil as an Ecosystem
Not to be forgotten in this mix are the microbes, insects, nematodes,
worms, and other organisms that, along with fungi and plants, occupy the
soil ecosystem. There are at least twelve different classifications of soil rec-
ognized by the U.S. Department of Agriculture, and these reflect the many
communities that occupy various soil ecosystems. In fact, when ecologists
discuss soil as an ecosystem, they are referring not only to the biotic and
abiotic components of soil itself but also to the living organisms, from all
kingdoms, that occupy, partially occupy, or travel through soil. Disruption
596
Soil
to soil by herbicides, polluted water, solid waste, mechanical erosion, and

other factors therefore has a reverberating impact not only on the minerals
and chemical compounds that form soil but also on all members of the soil
ecosystem.
Soil Chemistry
Soil ecologists and soil chemists are concerned not only with the composi-
tion of soil and soil water but also with how that composition changes as
the water interacts with the atmosphere, minerals, plants, fungi, animals,
and mechanical forces at work on it. Soil and its chemistry can be studied
in its natural environment, or samples can be brought into the laboratory
for testing. Some tests have been standardized and are best conducted in
the laboratory so that they can be compared with the results of other re-
searchers. Most of the standardized tests, such as measures of the soil’s
acidity and cation-exchange capacity, are related to measures of the soil’s
fertility and its overall suitability for plant growth. These tests measure av-
erage values for a soil sample because large original samples are dried and
thoroughly mixed before smaller samples are taken for the specific test.
Increasingly, soil chemists are looking for ways to study the fine details of
soil chemical processes. They know, for example, that soil water chemistry
changes as the water percolates through succeeding layers of the soil. The
water flowing through the soil during a rainstorm has a different chemical
composition from that of water clinging to soil particles, at the same depth,
several days later. Finally, during a rainstorm, the water flowing through
large cracks in the soil has a chemical composition different from that of the
same rainwater flowing through the tiny spaces between soil particles.
Sampling Techniques
Soil chemists use several sampling techniques to collect the different types
of soil water. During a rainstorm, water flows under the influence of grav-
ity. After digging a trench in the area of interest, researchers push several
sheets of metal or plastic, called pan lysimeters, into the wall of the trench
at specified depths below the surface. The pans have a very shallow V
shape. Soil water flowing through the soil collects in the pan, flows toward
the bottom of the V, and flows out of the pan into a collection bottle. Com-
paring the chemical compositions of rainwater that has passed through
different thicknesses of soils (marked by the depth of each pan) allows the
soil chemist to identify specific soil reactions with specific depths.
After the soil water stops flowing, water is still trapped in the soil. The
soil water clings to soil particles and is said to be held by tension. Tension
water can spend a long time in the soil between rainstorms. During that
597
The Twelve Soil Orders in the U.S. Classification System
Soil Order Features
Alfisols Soils in humid and subhumid climates with precipitation from 500 to
1,300 millimeters (20 to 50 inches), frequently under forest vegetation.
Clay accumulation in the B horizon and available water most of the
growing season. Slightly to moderately acid soils.
Andisols Soils with greater than 60 percent volcanic ash, cinders, pumice, and
basalt. They have a dark A horizon as well as high absoption and
immobilization of phosphorus and very high cation exchange capacity.
Aridisols Aridisols exist in dry climates. Some have horizons of lime or gypsum
accumulations, salty layers, and A and slight B horizon development.
Entisols Soils with no profile development except a shallow A horizon. Many
recent river floodplains, volcanic ash deposits, severely eroded areas,
and sand are entisols.
Gelisols Soils that commonly have a dark organic surface layer and mineral
layers underlain by permafrost, which forms a barrier to downward
movement of soil solution. Common in tundra regions of Alaska.
Alternate thawing and freezing of ice layers results in special features in
the soil; slow decomposition of the organic matter due to cold
temperatures results in a peat layer at the surface in many gelisols.
Histosols Organic soils of variable depths of accumulated plant remains in bogs,
marshes, and swamps.
Inceptisols Soils found in humid climates that have weak to moderate horizon
development. Horizon development may have been delayed because of
cold climate or waterlogging.
Mollisols Mostly grassland soils, but with some broadleaf forest-covered soils with
relatively deep, dark A horizons, a possible B horizon, and lime
accumulation.
Oxisols Excessively weathered soils. Oxisols are over 3 meters (10 feet) deep,
have low fertility, have dominantly iron and aluminum oxide clays, and
are acid. Oxisols are found in tropical and subtropical climates.
Spodosols Sandy leached soils of the cool coniferous forests, usually with an
organic or O horizon and a strongly acidic profile. The distinguishing
feature of spodosols is a B horizon with accumulated organic matter plus
iron and aluminum oxides.
Ultisols Strongly acid and severely weathered soils of tropical and subtropical
climates. They have clay accumulation in the B horizon.
Vertisols Soils with a high clay content that swell when wet and crack when dry.
Vertisols exist in temperate and tropical climates with distinct dry and
wet seasons. Usually vertisols have only a deep self-mixing A horizon.
When the topsoil is dry, it falls into the cracks, mixing the soil to the
depth of the cracks.
598
Soil
time, it reacts with soil mineral grains and soil microorganisms. Tension
water is sampled by placing another type of lysimeter, a tension lysimeter,
into the soil at a known depth. A tension lysimeter is like the nozzle of a
vaccum cleaner with a filter over the opening. Soil chemists actually vac-
uum the tension water out of the soil and to the surface for analysis.
Determining Isotopic Composition

Nonradioactive, stable isotopes of common elements are being used more
often by soil chemists to trace both the movement of water through the soil
and the chemical reactions that change the composition of the water. Trace
stable isotopes behave chemically just the way their more common coun-
terparts do. For example, deuterium, an isotope of hydrogen, substitutes
for hydrogen in the water molecule and allows the soil chemist to follow
the water’s movements. Similarly, carbon 13 and nitrogen 15 are relatively
rare isotopes of common elements that happen to be biologically impor-
tant. Using these isotopes, soil chemists can study the influences of soil or-
ganisms on the composition of soil water. Depending on what the soil
chemist is studying, the isotope may be added, or spiked, to the soil in the
laboratory or in the field. Alternatively, naturally occurring concentrations
of the isotope in rain or snowmelt may be used. Regardless, soil water sam-
ples are collected by one or more of the lysimeter methods, and their isoto-
pic composition is determined.
The Soil Chemical Factory

The wonderful interactions of complex chemical and biochemical reac-
tions that are soil chemistry are one indication of the uniqueness of planet
Earth. Without the interaction of liquid water and the gases in the atmo-
sphere, many of the nutrients necessary for life would remain locked up in
rock minerals. Thanks to weathering reaction, the soil chemical factory
started to produce nutrients, which resulted in the exploitation of the soil
environment by millions of organisms. The processes involved in soil
chemistry—from weathering reactions that turn rock into new soil to the
recycling of plant nutrients through microbial decay—are vital to every
human being. Without fertile soil, plants will not grow. Without plants as a
source of oxygen and food, there would be no animal life.
Because of the complex chemical interrelationships that have devel-
oped in the soil environment, it may seem that nothing can disrupt the
“factory” operation. As more is understood about soil chemistry and the
ways in which humans stress the soil chemistry through their activities, it
is apparent that the factory is fragile. Not only do humans rely on soil fer-
tility for their very existence, but they also are taking advantage of soil
599
Soil
chemical processes to help them survive their own past mistakes. Soil has
been and continues to be used as a garbage filter. Garbage, whether solid or
liquid, has been dumped on or buried in soil for ages. Natural chemical
processes broke down the garbage into simpler forms and recycled the nu-
trients. When garbage began to contain toxic chemicals, those chemicals,
when in small quantities, were either destroyed by soil bacteria or firmly
attached to soil particles. The result is that water—percolating through
garbage, on its way to the local groundwater, stream, or lake—does not
carry with it as much contamination as one might expect. Soil chemistry
has, so far, kept contaminated garbage from ruining drinking water. There
are well-known cases, however, where the volume and composition of
waste buried or spilled were such that the local soil chemistry was over-
whelmed. In cases of large industrial spills, or when artificial chemicals are
spilled or buried, the soil needs help to recover. The recovery efforts are
usually very expensive but, faced with the possible permanent loss of large
parts of the soil chemical factory, humankind cannot afford to neglect this
aspect of the environment.
Richard W. Arnseth
See also: Acid deposition; Deforestation; Endangered plant species; Ero-

sion and erosion control; Grasslands and prairies; Grazing and overgraz-
ing; Multiple-use approach; Nutrient cycles; Pesticides; Rangeland; Refor-
estation; Slash-and-burn agriculture; Soil contamination.

Berner, Elizabeth K., and Robert A. Berner. The Global Water Cycle: Geochem-
istry and Environment. Englewood Cliffs, N.J.: Prentice-Hall, 1987.
Bohn, Heinrich, B. L. McNeal, and G. A. O’Connor. Soil Chemistry. 2d ed.
New York: Wiley, 1985.
Brill, Winston. “Agricultural Microbiology.” Scientific American 245 (Sep-
tember, 1981): 198.
Evangelou, V. P. Environmental Soil and Water Chemistry: Principles and Ap-
plications. New York: Wiley, 1998.
Lloyd, G. B. Don’t Call It Dirt. Ontario, Calif.: Bookworm Publishing, 1976.
McBride, Murray B. Environmental Chemistry of Soils. New York: Oxford
Millot, Georges. “Clay.” Scientific American 240 (April, 1979): 108.
Sparks, Donald S., ed. Soil Physical Chemistry. 2d ed. Boca Raton, Fla.: CRC
Press, 1999.
Tan, Kim Howard. Principles of Soil Chemistry. 3d ed. New York: M. Dekker,
1998.
600
SOIL CONTAMINATION
Types of ecology: Ecotoxicology; Restoration and conservation ecology;
Soil ecology
Soils contaminated with high concentrations of hazardous substances pose poten-

tial risks to human health and the earth’s thin layer of productive soil.
P roductive soil depends on bacteria, fungi, and other soil microbes to

break down wastes and release and cycle nutrients that are essential to
plants. Healthy soil is essential for growing enough food for the world’s in-
creasing population. Soil also serves as both a filter and a buffer between
human activities and natural water resources, which ultimately serve as
the primary source of drinking water. Soil that is contaminated may serve
as a source of water pollution through leaching of contaminants into
groundwater and through runoff into surface waters such as lakes, rivers,
and streams.
Types of Contamination
Soils can become contaminated by many human activities, including fertil-
izer or pesticide application, direct discharge of pollutants at the soil surface,
leaking of underground storage tanks or pipes, leaching from landfills,
and atmospheric deposition. Additionally, soil contamination may be of nat-
ural origin. For example, soils with high concentrations of heavy metals can
occur naturally because of their close proximity to metal ore deposits. Com-
mon contaminants include inorganic compounds such as nitrate and heavy
metals (for example, lead, mercury, cadmium, arsenic, and chromium);
volatile hydrocarbons found in fuels, such as benzene, toluene, ethylene,
and xylene BTEX compounds; and chlorinated organic compounds such
as polychlorinated biphenyls (PCBs) and pentachlorophenol (PCP).
Contaminants may also include substances that occur naturally but
whose concentrations are elevated above normal levels. For example, ni-
trogen- and phosphorus-containing compounds are often added to agri-
cultural lands as fertilizers. Since nitrogen and phosphorus are typically
the limiting nutrients for plant and microbial growth, accumulation in the
soil is usually not a concern. The real concern is the leaching and runoff of
the nutrients into nearby water sources, which may lead to oxygen deple-
tion of lakes as a result of the eutrophication encouraged by those nutri-
ents. Furthermore, nitrate is a concern in drinking water because it poses a
direct risk to human infants; it is associated with blue-baby syndrome.
601
Soil contamination
Environmental Interactions
Contaminants may reside in the solid, liquid, and gaseous phases of the
soil. Most will occupy all three phases but will favor one phase over the
others. The physical and chemical properties of the contaminant and the
soil will determine which phase the contaminant favors. The substance
may preferentially adsorb to the solid phase, either the inorganic minerals
or the organic matter. The attraction to the solid phase may be weak or
strong. The contaminant may also volatize into the gaseous phase of the
soil. If the contaminant is soluble in water, it will dwell mainly in the liquid-
filled pores of the soil.
Contaminants may remain in the soil for years or make their way into
the atmosphere or nearby water sources. Additionally, the compounds
may be broken down or taken up by the biological component of the soil.
This may include plants, bacteria, fungi, and other soil-dwelling microbes.
The volatile compounds may slowly move from the gaseous phase of the
soil into the atmosphere. The contaminants that are bound to the solid
phase may remain intact or be carried off in runoff attached to soil particles
and flow into surface waters. Compounds that favor the liquid phase, such
as nitrate, will either move into surface waters or leach down into the
groundwater.
Metals display a range of behaviors. Some bind strongly to the solid
phase of the soil, while others easily dissolve and wind up in surface or
groundwater. PCBs and similar compounds bind strongly to the solid sur-
face and remain in the soil for years. These compounds can still pose a
threat to waterways because, over long periods of time, they slowly dis-
solve from the solid phase into the water at trace quantities. Fuel compo-
nents favor the gaseous phase but will bind to the solid phase and dissolve
at trace quantities into the water. However, even trace quantities of some
compounds can pose a serious ecological or health risk. When a contami-
nant causes a harmful effect, it is classified as a pollutant.
Treatments
There are two general approaches to cleaning up a contaminated soil site:
treatment of the soil in place (in situ) or removal of the contaminated soil
followed by treatment (non-in situ). In situ methods, which have the ad-
vantage of minimizing exposure pathways, include biodegradation, vola-
tilization, leaching, vitrification (glassification), and isolation or contain-
ment. Non-in situ methods generate additional concerns about exposure
during the process of transporting contaminated soil. Non-in situ options
include thermal treatment (incineration), land treatment, chemical extrac-
tion, solidification or stabilization, excavation, and asphalt incorporation.
602
Soil contamination
The choice of methodology will depend on the quantity and type of con-
taminants and on the nature of the soil. Some of these treatment technolo-
gies are still in the experimental phase.
John P. DiVincenzo
See also: Acid deposition; Biomagnification; Biopesticides; Erosion and

erosion control; Food chains and webs; Geochemical cycles; Hydrologic
cycle; Integrated pest management; Pesticides; Pollution effects; Soil;
Waste management.

Pierzynski, Gary M., J. Thomas Sims, and George F. Vance. Soils and Envi-
ronmental Quality. Boca Raton, Fla.: Lewis, 1994.
Sparks, Donald L. “Soil Decontamination.” In Handbook of Hazardous Mate-
rials, edited by Morton Corn. San Diego, Calif.: Academic Press, 1993.
Testa, Stephen M. The Reuse and Recycling of Contaminated Soil. Boca Raton,
Fla.: Lewis, 1997.
603
SPECIATION
Types of ecology: Community ecology; Evolutionary ecology; Speciation
Processes whereby new species arise is referred to as speciation. The term is used
most often to refer to the multiplication of species.
M any types of speciation have been proposed, but most can be grouped
into three main modes. Geographic (allopatric) speciation depends
upon geographic isolation of populations. Semigeographic (parapatric)
speciation involves divergence between populations in continuous geo-
graphic contact. Nongeographic (sympatric) speciation involves specia-
tion at a restricted locality, without geographic separation.
Allopatric Speciation
Geographic, or allopatric, speciation is widely accepted as the most impor-
tant mode of speciation for sexual species. According to this model, a new
species develops when a population becomes geographically isolated
from the remainder of the species and gradually evolves independently to
the extent that it becomes reproductively isolated. If the two populations
reestablish contact subsequent to the development of reproductive isola-
tion, no interbreeding will take place. Geographic speciation is a slow pro-
cess not amenable to experimental testing, but abundant indirect evidence
is furnished by patterns of geographic variation, the development of re-
productive isolation between geographically remote elements of the same
species, varying degrees of divergence between isolates, and correlation
of speciation with periods of isolation produced by past climatic or geolog-
ical events. An unproven variant of the allopatric model is the founder-
effect model, in which an isolated population established by a few found-
ers goes through a drastic genetic reorganization because of the limited
genetic variability introduced by the founders and chance fluctuations in
gene frequencies (genetic drift). Such a reorganization supposedly could
accelerate the speciation process.
Parapatric Speciation
The possibility of parapatric (semigeographic) speciation is suggested by
the occurrence of hybrid zones or belts along which two races interbreed
freely. Many hybrid belts are undoubtedly the result of secondary contact
between previously isolated populations, but others may have been
formed in place. The origin of hybrid zones, interactions at hybrid zones,
604
Speciation
and processes, if any, whereby discontinuities can proceed to reproductive

isolation are subjects of continuing investigation. Most chromosomal mod-
els are also essentially parapatric. Such models usually involve fixation in
a local population of chromosomal rearrangements that severely reduce
fertility in heterozygotes. Small, semi-isolated populations or mating sys-
tems that promote inbreeding are usually required. The subject is contro-
versial.
Sympatric Speciation
The origin of an asexual form from a sexual one always takes place in a
nongeographic (sympatric) mode. There are many variants of asexual repro-
duction, including egg or seed development without fertilization (parthe-
nogenesis and agamospermy, respectively), vegetative reproduction, and
simple fission. Obligatory self-fertilization in hermaphrodites accomplishes
the same end. Considerable attention has been given to the adaptive signif-
icance of uniparental reproduction and factors influencing its origin.
The only unquestioned mode of sympatric speciation for sexual forms
is through polyploidy, with doubling of the entire complement of chromo-
somes, a phenomenon often associated with hybridization. Tetraploids
(4N), for example, produced by a doubling of the diploid (2N) chromo-
some number, are often fertile, but are reproductively isolated from the pa-
rental forms because backcrossing produces sterile triploids (3N). Sexual
polyploids are known from many animal groups but are generally rare.
Polyploids are common among plants, however, and polyploidy has been
an important speciation mechanism in plants.
Many other models for sympatric speciation have been proposed, and
most are controversial. Many of these models involve disruptive selection,
a type of selection in which two or more phenotypes have high fitness,
while intermediates between them have low fitness. If an organism, for ex-
ample, exploits two subniches in the same locality, and there is some im-
pediment to free gene exchange, disruptive selection could produce two
different genotypes, each adapted to one of the subniches; heterozygotes
would be adapted to neither. This type of speciation has received some
support from studies on herbivorous insects and insects with parasitic lar-
vae; conditioning may favor egg deposition by the insect on the plant or
host where it developed. This would inhibit free gene exchange and allow
adaptation to different plants or hosts.
Ecological and Evolutionary Implications

The species is the most fundamental unit of classification and is the basic
unit of reference with respect to living things. It is an important unit of in-
605
Speciation
teraction in natural communities. It is implicit that a species is genetically

distinct from other species, with its own morphological and physiological
attributes; as such it is an essential reference in all fields of biology and in
the applied life sciences as well.
The origin of a new species signifies the appearance of a distinct evolu-
tionary unit with its own potential and that is isolated by intrinsic barriers
from other species.
The evolutionary implications for sexual and asexual forms, however,
are quite different. Sexual forms have an immeasurably greater potential
for evolutionary change because of the reservoir of genetic variability of
the population, which is shared and reshuffled through sexual reproduc-
tion. Asexual forms have a limited capacity for change, rarely give rise to
anything new, and tend to exploit short-term opportunities. The majority
of described species other than microorganisms are sexual, although many
exploit both modes of reproduction.
The speciation process itself does not necessarily produce adaptive
change, although changes in adaptation are usually involved in speciation.
Species, however, once formed, interact with other species through com-
petition, and selective forces promote specialization and adaptation to
subniches. This process leads not only to more diverse and complex eco-
logical communities but also to more pronounced morphological and
physiological differences between species. This phenomenon is well illus-
trated by various adaptive radiations that have taken place on initially
depauperate islands or in species-poor lakes following colonization by one
or a few species.
Speciation in this light can be seen to be a critical step that opens the
way to, and indirectly promotes, evolutionary change. The origin of higher
taxonomic categories has also been linked to speciation. The fossil record
indicates that higher categories originate when there is movement into a
major and/or distinctive and previously unexploited adaptive zone, fol-
lowed by further morphological change and diversification within the
zone. If a multiplicity of species are adapted to different subniches, chances
are greatly enhanced for the “discovery” of a new adaptive zone by at least
one species. Once movement into the zone is accomplished, diversification
through speciation would take place, and interactions between species
would promote further change.
John S. Mecham
See also: Adaptations and their mechanisms; Adaptive radiation; Clines,

hybrid zones, and introgression; Convergence and divergence; Develop-
ment and ecological strategies; Evolution: definition and theories; Evolu-
606
Speciation
tion: history; Evolution of plants and climates; Extinctions and evolution-

ary explosions; Gene flow; Genetic drift; Isolating mechanisms; Natural
selection; Nonrandom mating, genetic drift, and mutation; Population ge-
netics; Punctuated equilibrium vs. gradualism; Species loss; Succession.

Claridge, Michael F., H. A. Dawah, and M. R. Wilson, eds. Species: The Units
of Biodiversity. New York: Chapman and Hall, 1997.
Lambert, David M., and Hamish G. Spencer, eds. Speciation and the Recogni-
tion Concept: Theory and Application. Baltimore: Johns Hopkins Univer-
sity Press, 1995.
Mayr, Ernst. Populations, Species, and Evolution. Cambridge, Mass.: Belknap
Press, 1970.
Otte, Daniel, and John Endler, eds. Speciation and Its Consequences. Sunder-
land, Mass.: Sinauer Associates, 1989.
Paterson, H. E. H. Evolution and the Recognition Concept of Species: Collected
Writings. Baltimore: Johns Hopkins University Press, 1993.
Townsend, Colin R., et al. Essentials of Ecology. 2d ed. Malden, Mass.:
Blackwell Science, 2003.
White, Michael. Modes of Speciation. San Francisco: W. H. Freeman, 1978.
607
SPECIES LOSS
Types of ecology: Evolutionary ecology; Restoration and conservation
ecology
Species loss, particularly the extinction of species that is caused by human activi-
ties, has increasingly concerned scientists in a number of fields, threatening biodi-
versity both locally and globally.
P ublic and scientific concern about species loss stems from several fac-
tors and encompasses a variety of viewpoints. Ethically, many people
believe that species have value in and of themselves and that humankind
does not have the right to cause the extinction of any species. A species
may also have an unknown potential to enrich human life and health. The
latter argument is important in that many synthetic medicines and com-
mercial products were first produced by plants and animals. The loss of
species could potentially mean the loss of beneficial new products for hu-
man society. Species that exist today are the result of millions of years of
evolutionary success, and to lose species is to lose that evolutionary his-
tory. From a resource management point of view, ecologists and land man-
agers alike are concerned about the effects that species loss may have on
the function and stability of biotic communities.
Ecological Concerns
Relationships such as predation, competition, and parasitism link species
into complex community relationships. One way species are linked is by
trophic levels within the community food chain, which is more accurately
described as a food web. Starting with plants at the base of the web, trophic
levels begin with producers, followed by several successive levels of con-
sumers: herbivore, first-level carnivore, second-level carnivore, and so on,
up to top carnivore. Omnivores feed both as herbivores and as carnivores
and thus feed at more than one trophic level. Finally decomposers feed on
dead organisms and their waste products from all trophic levels.
Therefore, although the ramifications of loss of a species are not easily
predicted, such a loss will have significant impact on ecosystems—often
beyond merely the obvious one of increasing a population of its prey or de-
creasing a population of predators. Such a loss will disturb the entire food
chain—or, more properly, food web—involving the complex relationships
among all species in a community. Such community disturbance inevitably
expands beyond the community’s borders to affect the larger ecosystem.
608
Species loss
Some examples of these complex relationships have been revealed by con-

trolled studies.
Species-Removal Studies
Species-removal studies provide some indication of what may occur when
a species becomes extinct. In more than 90 percent of predator-removal
studies, population densities of prey species in the trophic level immedi-
ately below the predator have shown a significant increase or decrease. In
many cases, the change in density was twofold. Rarely has the removal of
predator species had no effect on the population density of its prey. How-
ever, not all studies have shown the expected increase in prey density;
many have shown an unexpected decrease.
For species that possibly compete with one another, more than 90 per-
cent of competitor-removal studies have shown an increase in the “remain-
ing competitor” population density. Several factors may influence the
strength of community response in species-removal experiments. For ex-
ample, a predator may prey more heavily on a large, aggressive prey spe-
cies and thus allow the coexistence of a less aggressive, competitor prey
species. If the predator is removed, the aggressive prey may increase in
density while the less aggressive one may actually decrease. Studies in
aquatic communities indicate that the higher the trophic level in which
species removal occurs, the greater the effect on population densities at
lower trophic levels.
The ramifications of species loss can only partially be predicted with
knowledge of community food webs. The size and direction of population
density change within a community may or may not be as expected. It is
safe to predict, however, that species loss will cause changes in most in-
stances.
Wildlife Protection and Endangered-Species Legislation

Concern about species loss in North America can be traced back at least as
far as 1872, when legislation offering limited protection to the American bi-
son (buffalo) was passed by the United States Congress. This legislation
was passed at the height of buffalo exploitation by market hunters and
during the United States Army’s policy of fighting Native American tribes
by cutting off their food supply. However, President Ulysses S. Grant ve-
toed the legislation, and the buffalo was almost lost. Only a few hundred
remained by 1900. The first National Wildlife Refuge was set aside by Pres-
ident Theodore Roosevelt in 1902 to protect egrets from extinction by
feather hunters. Three years later, the Wichita Mountain National Wildlife
Refuge was set aside to protect one of the small remnant herds of buffalo.
609
Species loss
Several North American species and subspecies are now extinct because of
similar exploitations: The passenger pigeon, Carolina parakeet, heath hen,
Merriam’s elk, and Badlands bighorn sheep are some of the best known ex-
amples.
During the 1960’s increasing concern about an accelerated species ex-
tinction rate attributable to human exploitation and disturbance of the
environment culminated in the first federal protective legislation for en-
dangered species, the Endangered Species Preservation Act of 1966. This
act was limited to listing endangered birds and mammals and funding
research on their population ecology and habitat acquisition. This legis-
lation was expanded in 1969 to include all vertebrate animal species
and some invertebrates. The definitive protection legislation is the 1973
Endangered Species Act. This act set procedures for listing threatened
and endangered species, called for designation of critical habitat for each
threatened or endangered species, and mandated the development of
recovery plans for these species. The act prohibits the use of federal funds
for projects that would harm threatened or endangered species. The cover-
age of the 1973 act was also expanded to include plants and invertebrate
animals (except pest insects), subspecies, and distinct vertebrate popula-
tions.
Since 1966 the U.S. Fish and Wildlife Service (USFWS) has had the legal
responsibility of compiling and maintaining an official threatened and
endangered species list. There are formal petitioning processes for plac-
ing additional species on the list and for removing them from the list. Peti-
tions may be initiated by the USFWS or by private organizations. Petitions
are reviewed by scientific panels using all available information on the
species. If sufficient information is available to support the petition, a pro-
posed addition to the list is published in the Federal Register and other
appropriate places to solicit public comment. Final decisions about list-
ing, “down-listing” (for example, changing a species designation from
“endangered” to “threatened”), or “de-listing” are made by the USFWS.
The ultimate goal of the listing process and the implementation of a recov-
ery plan is to increase the abundance and distribution of a species to the
point of being able to remove it from the threatened and endangered spe-
cies list.
James F. Fowler
See also: Biodiversity; Conservation biology; Deforestation; Endangered

animal species; Endangered plant species; Extinctions and evolutionary
explosions; Habitats and biomes; Old-growth forests; Reforestation; Resto-
ration ecology; Speciation; Wildlife management; Zoos.
610
Species loss

Sherry, Clifford J. Endangered Species: A Reference Handbook. Santa Barbara,
Calif.: ABC-Clio, 1998.
Stanley, Steven M. Extinction. New York: Scientific American Library, 1987.
Stearns, Beverly Petersen, and Stephen C. Stearns. Watching, from the Edge
of Extinction. New Haven, Conn.: Yale University Press, 1999.
Ward, Peter D., and Don Brownlee. Rare Earth: Why Complex Life Is Uncom-
mon in the Universe. New York: Copernicus, 2000.
Wilson, Edward O. The Diversity of Life. New York: W. W. Norton, 1993.
_______. The Future of Life. New York: Alfred A. Knopf, 2001.
611
SUCCESSION
Succession is the progressive and orderly replacement of one biological community

by another until a relatively stable, self-maintaining “climax community” is
achieved.
S uccession is an important ecological phenomenon because it allows the

maximum variety and number of species to occupy a given area
through time and leads to the establishment of an ecologically stable cli-
max community that represents the most complex and diverse biological
system possible, given existing environmental conditions and available
energy input. As succession proceeds, significant changes occur in species
composition, nutrient cycling, energy flow, productivity, and stratification.
Changes also occur within the climax community; however, these changes
act to maintain the climax, not alter it.
Immature communities tend to have high populations of a few species
that are relatively small and simple. Biomass (weight of living material) is
low, and nutrient conservation and retention are poor. Food chains are
short, and available energy is shared by few species. Community structure
is simple and easily disrupted by external forces. As communities mature,
larger and more complex organisms appear, and there is a higher species
diversity (number of different species). Biomass increases, and nutrients
are retained and cycled within the community. The greater number of spe-
cies results in more species interactions and the development of complex
food webs. Community productivity (conversion of solar energy to chemi-
cal energy), initially high in immature communities, becomes balanced by
community respiration as more energy is expended in maintenance activi-
ties.
Stages of Succession
The entire sequence of communities is called a sere, and each step or com-
munity in the sequence is a seral stage. The climax community is in bal-
ance, or equilibrium, with the environment and displays greater stability,
more efficient nutrient and energy recycling, a greater number of species,
and a more complex community structure than that of each preceding seral
stage.
Each seral stage is characterized by its own distinctive forms of plant
and animal life, which are adapted to a unique set of chemical, physical,
612
Succession
and biological conditions. Excepting the climax community, change is the

one constant shared by all seral stages. Changes can be induced by abiotic
factors, such as erosion or deposition, and by biotic factors, modification of
the environment caused by the activities of living organisms within the
community.
These self-induced factors bring about environmental changes detri-
mental to the existing community but conducive to invasion and replace-
ment by more suitably adapted species. For example, lichens are one of the
first colonizers of barren rock outcrops. Their presence acts to trap and
hold windblown and water-carried debris, thereby building up a thin soil.
As soil depth increases, soil moisture and nutrient content become optimal
for supporting mosses, herbs, and grasses, which replace the lichens.
These species continue the process of soil-building and create an environ-
ment suitable for woody shrubs and trees.
In time, the trees overtop the shrubs and establish a young forest. These
first trees are usually shade-intolerant species. Beneath them, the seeds of
the shade-tolerant trees germinate and grow up, eventually replacing the
shade-tolerant species. Finally, a climax forest community develops on
what once was bare rock, and succession ends.
Primary and Secondary Succession

The sere just described—from barren rock to climax forest—is an example
of primary succession. In primary succession, the initial seral stage, or pio-
neer community, begins on a substrate devoid of life or unaltered by living
organisms. Succession that starts in areas where an established community
has been disturbed or destroyed by natural forces or by human activities
(such as floods, windstorms, fire, logging, and farming) is called second-
ary succession.
An example of secondary succession occurs on abandoned cropland.
This is referred to as old-field succession and begins with the invasion of
the abandoned field by annual herbs such as ragweed and crabgrass.
These are replaced after one or two years by a mixture of biennial and pe-
rennial herbs, and by the third year the perennials dominate. Woody
shrubs and trees normally replace the perennials within ten years. After
another ten or twenty years have passed, a forest is established, and ulti-
mately, after one or two additional seral stages in which one tree commu-
nity replaces another, a climax forest emerges.
Both primary and secondary succession begin on sites typically low in
nutrients and exposed to extremes in moisture, light intensity, tempera-
ture, and other environmental factors. Plants colonizing such sites are tol-
erant of harsh conditions, are characteristically low-growing and relatively
613
Succession
small, and have short life cycles. By moderating the environmental condi-
tions, these species make the area less favorable for themselves and more
favorable for plants that are better adapted to the new environment. Such
plants are normally long-lived and relatively large. Secondary succession
usually proceeds at a faster rate than primary succession, because a well-
developed soil and some life are already present.
Aquatic Environments
Succession can also take place in aquatic environments, such as a newly
formed pond. The pioneer community consists of microscopic organisms
that live in the open water. Upon death, their remains settle on the bottom
and join with sediment and organic matter washed into the pond. An accu-
mulation of sediment provides anchorage and nutrients for rooted, sub-
merged aquatic plants such as pondweeds and waterweeds. These add to
the buildup of sediment, and as water depth decreases, rooted, floating-
leaved species such as water lilies prevent light from reaching the sub-
merged aquatics and eliminate them.
At the water’s edge, emergent plants rooted in the bottom and extend-
ing their stems and leaves above water (cattails, rushes, and sedges) trap
Image Not Available
614
Succession
sediment, add organic matter, and continue the filling-in process. The shal-
low margins fill first, and eventually the open water disappears and a
marsh or bog forms. A soil rich in partially decomposed organic matter
and saturated with water accumulates. As drainage improves and the soil
becomes raised above the water level, trees and shrubs tolerant of wet soils
invade the marsh. These act to lower the water table and improve soil aera-
tion. Trees suited to drier conditions move in, and once again a climax com-
munity characteristic of the surrounding area develops.
Theories of Succession
The American ecologist Frederic E. Clements (1874-1945) believed that the
characteristics of a climax community were determined solely by regional
climate. According to Clements, all communities within a given climatic
region, despite initial differences, eventually develop into the same climax
community. Some seral stages might be abbreviated or skipped entirely,
while others could be lengthened or otherwise modified; however, the end
result would always be a single climax community suited to the regional
climate. This phenomenon is called convergence, and Clements’s single-
climax concept is known as the monoclimax theory.
Some ecologists have found the monoclimax theory to be simplistic and
have offered other theories. One of these, the polyclimax theory, holds that,
within a given climatic region, there could be many climaxes. It was noted
that in any single climatic region, there were often many indefinitely main-
tained communities that could be considered separate and distinct cli-
maxes. These developed as a result of differences caused by soil type, soil
moisture, nutrients, slope, fire, animal activity (grazing and browsing),
and other factors. Clements countered that these would eventually reach
true climax status if given enough time and proposed terms such as
subclimax (a long-lasting seral stage preceding the climax) and disclimax
(a nonclimax maintained by continual disturbance) to describe such situa-
tions.
A third theory, the climax pattern concept, views the climax as a single
large community composed of a mosaic or pattern of climax vegetation in-
stead of many separate climaxes or subclimaxes. Numerous habitat and
environmental differences account for the patterns of populations within
the climax; no single factor such as climate is responsible.
While there is little doubt about the reality of succession, it is apparently
not a universal phenomenon. For example, disturbed areas within tropical
rain forests do not undergo a series of seral stages leading to reestablish-
ment of the climax community. Instead, the climax is established directly
by the existing species. Nevertheless, in most regions succession is the
615
Succession
mechanism by which highly organized, self-maintained, and ecologically

efficient communities are established.
Basis for Biodiversity

Succession is an important ecological phenomenon because it allows the
maximum variety and number of species to occupy a given area through
time and leads to the establishment of an ecologically stable climax com-
munity that represents the most complex and diverse biological system
possible, given existing environmental conditions and available energy in-
put. As succession proceeds, significant changes occur in species composi-
tion, nutrient cycling, energy flow, productivity, and stratification. Changes
also occur within the climax community; however, these changes act to
maintain the climax, not alter it.
Immature communities tend to have high populations of a few species
that are relatively small and simple. Biomass (weight of living material) is
low, and nutrient conservation and retention are poor. Food chains are
short, and available energy is shared by few species. Community structure
is simple and easily disrupted by external forces. As communities mature,
larger and more complex organisms appear, and there is a higher species
diversity (number of different species). Biomass increases, and nutrients
are retained and cycled within the community. The greater number of spe-
cies results in more species interactions and the development of complex
food webs. Community productivity (energy storage), initially high in im-
mature communities, becomes balanced by community respiration as
more energy is expended in maintenance activities. Community structure
increases in complexity and stability as equilibrium with the prevailing
physical environment is achieved.
Human beings, throughout their history, have been interacting with
natural communities and the process of succession—sometimes with di-
sastrous results. Much of what was once climax forest or grassland has
been put to the plow, timbered, strip-mined, or otherwise altered. In such
cases, humans retard or reverse succession by destroying or disrupting the
existing climax and replacing it with an ecologically simpler and less stable
seral stage. In a cornfield, for example, the existing climax has been re-
placed, in effect, by a simple pioneer community whose dominant species
is an annual. Invading weeds and shrubs must be constantly controlled or
eliminated. Nutrients, in the form of fertilizer, must be applied to maintain
high yields. Windstorms, drought, insect attacks, and other natural calami-
ties can easily destroy the entire community. A cornfield is neither self-
repairing nor self-maintaining, and humans must constantly intercede to
keep succession in check.
616
Succession
Although such human intervention is easily justified by its benefits, hu-

man exploitation of the natural environment is too often destabilizing and
destructive. The Dust Bowl of the 1930’s is an excellent example. The
shortgrass climax community that existed in parts of the southwestern
Great Plains was converted to wheat production and ranchland without
regard for the consequences. Drought, overgrazing, and poor farming
practices combined to convert the once-fertile prairie into a barren waste-
land. Failure to understand the physical and climatic conditions that re-
sulted in shortgrass prairie, coupled with unwise land usage, was the un-
derlying cause of this environmental disaster.
Steven D. Carey
See also: Biodiversity; Biogeography; Biomes: determinants; Biomes:

types; Coevolution; Communities: ecosystem interactions; Communities:
structure; Competition; Ecology: definition; Food chains and webs; Gene
flow; Genetic diversity; Speciation; Symbiosis; Trophic levels and ecologi-
cal niches.

Bazzaz, F. A. Plants in Changing Environments: Linking Physiological, Popula-
tion, and Community Ecology. New York: Cambridge University Press,
1996.
Brewer, Richard. The Science of Ecology. Fort Worth, Tex.: Saunders College
Publishers, 1994.
Perry, David A. Forest Ecosystems. Baltimore: Johns Hopkins University
Press, 1994.
Smith, Robert L. Ecology and Field Biology. 6th ed. San Francisco: Benjamin
Cummings, 2001.
617
SUSTAINABLE DEVELOPMENT
Types of ecology: Restoration and conservation ecology; Theoretical
ecology
Sustainable development meets the consumption needs of the current generation

without compromising the ability of future generations to increase their economic
production to meet future needs. Environmental benefits arise as a consequence of
changes in human attitude and behavior, technology, and resource utilization.
I n 1987, the United Nations World Commission on Environment and De-

velopment, also known as the Brundtland Commission, issued a report
in which it noted that humanity has the ability to make development “sus-
tainable”—to ensure that it meets the needs of the present without com-
promising the ability of future generations to meet their own needs. Sus-
tainable development is a process of change in which the exploitation of
resources, the direction of investments, the orientation of technological de-
velopment, and institutional change are all in harmony and enhance both
current and future potential to meet human needs and aspirations. The
commission envisioned the possibility of continued economic growth,
population stabilization, improvements in global economic equity be-
tween rich and poor nations, and environmental improvement, all occur-
ring simultaneously and in harmony. Since publication of the Brundtland
Commission’s report, sustainable development has become the dominant
global position on the environment, ecology, and economic development.
A Value System
Sustainable development is a normative philosophy, or value system, con-
cerned with equal distribution of the earth’s natural capital among current
and future generations of humans. Sustainable development promotes
three core values. First, current and future generations should each have
equal access to the planet’s life-support systems—including Earth’s gas-
eous atmosphere, biodiversity, stocks of exhaustible resources, and stocks
of renewable resources—and should maintain the earth’s atmosphere,
land, and biodiversity for future generations. Exhaustible resources, such
as minerals and fossil fuels, are used sparingly and conserved for use by
future generations. Renewable resources, such as forests and soil fertility,
are renewed as they are used to ensure that stocks are maintained at or
above current levels and are never exhausted.
Second, all future generations should have an equal opportunity to en-
618
Sustainable development
joy a material standard of living equivalent to that of the current genera-

tion. In addition, the descendants of the current generation in underdevel-
oped regions are permitted to increase their economic development to
match that available to descendants of the current generation in the indus-
trialized regions. Future development and growth in both developed and
underdeveloped regions must be sustainable.
Finally, future development must no longer follow the growth path
taken by the currently industrialized countries but should utilize appro-
priate technology. Development should also limit use of renewable re-
sources to each resource’s maximum sustained yield, the rate of harvest of
natural resources such as fisheries and timber that can be maintained in-
definitely through active human management of those resources.
Weak sustainability requires that depletions in natural capital be com-
pensated for by increases in human-made capital of equal value. For exam-
ple, the requirements for weak sustainability are met when a tree (natural
capital) is cut for the construction of a frame house (human-made capital).
However, if the tree is cut and cast aside in a land-clearing project, the re-
quirements for weak sustainability are not met. Strong sustainability re-
quires that depletions of one sort of natural capital be compensated for by
increases in the same or similar natural capital. For example, the require-
ments for strong sustainability are met when a tree is cut and a new tree is
planted to replace it, or when loss of acreage in equatorial rain forests in
Brazil is compensated for by an increase in the acreage of temperate rain
forests on the Pacific coast of North America.
Promoting the Philosophy

Sustainable development is promoted through a combination of public
policies. First, to the extent possible, elements in the earth’s support system
are assigned monetary values in order to make the economic and financial
calculations that are necessary to ensure that the requirements of weak
sustainability are met.
Second, economic development in the underdeveloped world is shifted
away from high-resource-using, high-polluting patterns of Western devel-
opment and toward more sustainable or “appropriate” patterns. Sug-
gested appropriate technologies include solar energy, resource recycling,
cottage industry, and microenterprises (factories built on a small scale).
Third, objective and measurable air, water, and resource quality stan-
dards are established and enforced to ensure that a continuing minimum
quality and quantity of natural capital is maintained and that certain
stocks of natural capital are protected through the establishment of wilder-
ness areas, oil and gas reserves, and other reserves.
619
Sustainable development
Finally, each individual human adopts a personal commitment to a sus-

tainable lifestyle, thus making a minimal personal impact on the earth’s
natural capital.
Environmental improvement results from the changes in resource utili-
zation. For example, reductions in use and waste of natural capital reduces
the environmental impact of resource extraction industries such as strip
mines, and waste disposal industries such as incinerators. Environmental
quality standards and maintenance of biodiversity leads to implementa-
tion of antipollution and ecosystem restoration efforts.
Gordon Neal Diem

animal species; Endangered plant species; Erosion and erosion control;
Forest management; Grazing and overgrazing; Integrated pest manage-
ment; Multiple-use approach; Old-growth forests; Reforestation; Restora-
tion ecology; Species loss; Urban and suburban wildlife; Waste manage-
ment; Wildlife management; Zoos.

Bowers, John. Sustainability and Environmental Economics: An Alternative
Text. Harlow, England: Longman, 1997.
Dryzek, John, ed. Debating the Earth: The Environmental Politics Reader. New
Lee, Kai N. Compass and Gyroscope: Integrating Science and Politics for the En-
vironment. Washington, D.C.: Island Press, 1993.
Sitarz, Daniel. Sustainable America: America’s Environment, Economy, and So-
ciety in the Twenty-first Century. Foreword by Al Gore. Carbondale, Ill.:
EarthPress, 1998.
United Nations Earth Summit. Agenda 21. New York: Author, 1992.
World Bank. Monitoring Environmental Progress. Washington, D.C.: Envi-
ronmentally Sustainable Development, World Bank, 1995.
_______. The World Bank and the Global Environment: A Progress Report.
Washington, D.C.: Author, 2000.
620
SYMBIOSIS
All animals live in close association, or symbiosis, with other species. Most symbi-
oses are based on nutritional interrelationships involving competition or coopera-
tion. Some animals cannot survive without their symbiotic partners, while others
are harmed or killed by them.
U nderstanding the ways in which different species of animals interact

in nature is one of the fundamental goals of ecology. Predator-prey re-
lationships, competition between species for limited resources, and symbi-
osis are the major forms of species interactions, and these have profoundly
influenced the diversity and ecology of all forms of life. Significant ad-
vances have been made in understanding how organisms interact, but in
studies of symbiosis (which literally means “living together”), one finds
the most complex, interesting, and important examples of both coopera-
tion and exploitation known in the living world.
Defining Symbiosis
Symbiosis involves many types of dependent or interdependent associa-
tions between species. In contrast to predator-prey interactions, however,
symbioses are seldom rapidly fatal to either of the associating species
(symbionts) and are often of long duration. With the exception of grazing
animals that do not often entirely consume or destroy their plant “prey,”
most predators quickly kill and consume their prey. While a predator may
share its prey with other individuals of the same species (clearly an exam-
ple of “living together”), such intraspecific behavior is not considered to be
a type of symbiosis. Fleas, some ticks, mites, mosquitoes, and other blood-
sucking flies are viewed as micropredators rather than parasites.
All organisms are involved in some form of competition. The abun-
dance and availability of environmental resources are finite, and competi-
tion for resources occurs both between members of the same species and
between individuals and populations of different species. When the num-
ber of individuals in a population increases, the intensity of competition
for limited food, water, shelter, space, and other resources necessary for
survival and reproduction also increases. Thus, competition plays a major
role in populations of free-living animals (those not inhabiting the body of
other organisms) and in populations living on or in other animals. For ex-
ample, both tapeworms and whales must compete for resources, and both
621
Symbiosis
have evolved habitat-specific adaptations to accomplish this goal. Whales

compete with whales, fish, and other predators for food; tapeworms com-
pete with tapeworms and other symbionts (such as roundworms) for food
and space; and tapeworms and whales compete with each other for food in
the whale’s gut.
“Symbiosis” is a term used to describe nonaccidental, nonpredatory as-
sociations between species. When used by itself, the term “symbiosis”
does not provide information on how or why species live together, or the
biological consequences of their interactions. Recognizably different forms
of symbioses all have one or more characteristics in common. All involve
“living together”; most involve food sharing; many involve shelter; and
some involve damage to one or both symbionts.
Hosts and Symbionts

Host species may be thought of as landlords. Hosts provide their symbi-
onts (also called symbiotes) with transportation, shelter, protection, space,
some form of nutrition, or some combination of these. Host species are
generally larger and structurally more complex than their symbionts, and
different parts of a host’s body (skin, gills, and gut, for example) may pro-
vide habitats for several different kinds of symbionts at the same time. The
three primary categories of symbiosis most commonly referred to in popu-
lar and scientific works are commensalism, mutualism, and parasitism.
Symbionts that share a common food source are known as commensals
(literally, “mess-mates”). In the usual definition of commensalism, one
species (usually referred to as the commensal, although both species are
commensals) is said to benefit from the relationship, while the other (usu-
ally referred to as the host) neither benefits nor is harmed by the other.
Adult tapeworms which live in the intestinal tracts of vertebrate hosts pro-
vide a classic example of commensals. Adult tapeworms share the host’s
food, usually with little or no effect on otherwise healthy hosts. As in all
species, however, too large a tapeworm population may result in excessive
competition, lower fitness, or disease in both the host and the tapeworms.
For example, the broad fish tapeworm, which includes humans among its
hosts, Diphyllobothrium latum, may cause a vitamin B-12 deficiency and
anemia in humans when the worm burden is high. In addition to tape-
worms, many human symbionts called “parasites” are, in fact, commen-
sals.
External commensals (those living on the skin, fur, scales, or feathers of
their hosts) are called epizoites. A good example of an epizoite is the fish
louse (a distant relative of the copepod), which feeds on mucus of the skin
and scales of fish. Another type of commensalism is called phoresis
622
Symbiosis
Remoras attach themselves to larger fish for transport. This form of symbiosis is
called phoresis, the passive transportation of the commensal (here, the remora) by
its host. (Digital Stock)
(phoresy), which involves passive transportation of the commensal

(phoront) by its host. Examples of phoreses include barnacles carried by
whales and sea turtles, and remoras (sharksuckers), which, in the absence
of sharks, may temporarily attach themselves to human swimmers. In
inquilinism, the transported commensal (inquiline) shares, or more accu-
rately, steals, food from the host, or may even eat parts of the host. Perhaps
the best-known inquilines are the glass- or pearlfish, which take refuge in
the cloacae of sea cucumbers and often eat part of the host’s respiratory
system. A unique type of commensalism, known as symphilism, is found
in certain ants and some other insects (hosts) which “farm” aphids (sym-
philes) and induce them to secrete a sugary substance which the ants eat.
Mutualism
The most diverse type of commensalism is mutualism. In some works, par-
ticularly those dealing with animal behavior, mutualism is used as a syn-
onym of symbiosis; hence, the reader must use caution in order to deter-
mine an author’s usage of these terms. As used here, mutualism is a special
case of commensalism, a category of symbiosis. The relationship between
mutuals may be obligatory on the part of one or both species, but it is al-
ways reciprocally beneficial, as the following examples illustrate.
623
Symbiosis
Some species of hermit crab place sea anemones on their shells or claws
(sea anemones are carnivores which possess stinging cells in their ten-
tacles). Hermit crabs without anemones on their shells or claws may be
more vulnerable to predators than those with an anemone partner. Hermit
crabs, which shred their food in processing it, lose some of the scraps to the
water, which the anemones intercept, and eat. Thus, the crab provides food
to the anemone, which in turn protects its provider. Such relationships,
which are species-specific, are probably the result of a long period of co-
evolution.
A different type of mutualism, but one having the same outcome as the
crab-anemone example, is found in associations between certain clown
fish and sea anemones. Clown fish appear to be fearless and vigorously at-
tack intruders of any size (including scuba divers) that venture too close to
“their” anemone. When threatened or attacked by predators, these small
fish dive into an anemone’s stinging tentacles, where they find relative
safety. Anemones apparently share in food captured by clown fish, which
have been observed to drop food on their host anemone’s tentacles.
Cleaning symbiosis is another unique type of mutualism found in the
marine environment. In this type of association, marine fish and shrimp of
several species “advertise” their presence by bright and distinctive color
patterns or by conspicuous movements. Locations where this behavior oc-
curs are called “cleaning stations.” Instead of being consumed by preda-
tory fish, these carnivores approach the cleaner fish or shrimp, stop swim-
ming, and sometimes assume unusual postures. Barracudas, groupers,
and other predators often open their mouths and gill covers to permit the
cleaners easy entrance and access to the teeth and gills. Cleaners feed on
epizoites, ectoparasites, and necrotic (dead) tissue that they find on host
fish, to the benefit of both species. Some studies have shown that removal
of cleaning symbionts from a coral reef results in a significant decrease in
the health of resident fish.
Parasitism
Parasitism is a category of symbiosis involving species associations that
are very intimate and in which competitive interactions for resources may
be both acute and costly. The extreme intimacy (rather than damage) be-
tween host and parasite is the chief difference between parasitism and
other forms of symbiosis. Parasites often, but not always, live within the
cells and tissues of their hosts, using them as a source of food. Some types
of commensals also consume host tissue, but in such cases (pearl fish and
sea cucumbers, for example) significant damage to the host rarely occurs.
Commensalism is associated with nutritional theft.
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Symbiosis
Some, but not all, parasites harm their hosts, by tissue destruction (con-
sumption or mechanical damage) or toxic metabolic by-products (ammo-
nia, for example). Commonly, however, damage to the host is primarily the
result of the host’s own immune response to the presence of the parasite in
its body, cells, or tissues. In extreme cases, parasites may directly or indi-
rectly cause the host’s death. When the host dies, its parasites usually die
as well. It follows that the vast majority of host-parasite relationships are
sublethal. A number of parasites are actually beneficial or crucial to the
survival of their hosts. The modern, and biologically reasonable, definition
of parasitism as an intimate type of symbiosis, rather than an exclusively
pathogenic association between species, promotes an ecological-evolu-
tionary understanding of interspecies associations. Most nonmedical ecol-
ogists and symbiotologists agree that two distinct forms of intimate associ-
ations, or parasitisms (with many intermediate types) occur in nature. The
most familiar are those involving decreased fitness in humans and in their
domestic animals and crops.
Among animal parasites, malarial parasites, hookworms, trypano-
somes, and schistosomes (blood flukes) cause death and disease in mil-
lions of people each year. The degree to which these parasites are patho-
genic, however, is partly the result of preexisting conditions of ill health,
malnutrition, other diseases, unsanitary living conditions, overcrowding,
or lack of education and prevention. Parasites which frequently kill or pre-
vent reproduction of their hosts do not survive in an evolutionary sense,
because both the parasites and their hosts perish. Both members of inti-
mate symbiotic relationships constantly adapt to their environments, and
to each other. Over time, evolutionary selection pressures result in co-
adaptation (lessening of pathogenicity) or destruction or change in form of
the symbiosis.
Nonpathogenic or beneficial host-parasite associations are among the
most highly evolved of reciprocal interactions between species. The ex-
treme degree of intimacy of the symbionts (not lack of pathogenicity) dis-
tinguishes this type of parasitism from mutualism. Parasitic dinoflagel-
lates (relatives of the algae that cause “red tides”) are found in the tissues
of all reef-building corals. These photosynthetic organisms use carbon
dioxide and other waste products produced by corals. In turn, the dino-
flagellates (Symbiodinium microadriaticum) provide their hosts with oxy-
gen and nutrients that the corals cannot obtain or produce by them-
selves. Without parasitic dinoflagellates, reef-building corals starve to
death. Similar host-parasite relationships occur in termites, which, with-
out cellulose-digesting parasitic protozoans in their gut, would starve to
death.
625
Symbiosis
Research on Commensals and Parasites

The life cycles of many commensals and parasites are extremely complex
and often involve two or more intermediate hosts living in different envi-
ronments, as well as free-living developmental stages. Knowledge of life
cycles remains as one of the most important areas of research in parasitol-
ogy and is usually the phase of research following the description of a new
species.
Scientists have long recognized that “chemical warfare” (antibiotics,
antihelminthics, insecticides) against microbial and animal parasites, and
their insect and other vectors, provides only short-term solutions to the
control or eradication of symbionts of medical importance. Research at-
tempts are being made to find ways of interrupting life cycles, sometimes
with the use of other parasites. This research requires sophisticated ecolog-
ical and biochemical knowledge of both the host-parasite relationship
and the parasite-mix. Studies of the parasite-mix are ecological (parasite-
parasite and host-parasite competition), immunological (host defense
mechanisms and parasite avoidance strategies), and ethological (host and
symbiont behavioral interactions) in nature. Investigators involved in this
kind of research must be well trained in many of the biological disciplines,
including epidemiology (the distribution and demographics of disease).
Immunology is the most promising modern research area in parasitol-
ogy. Not only have specific diagnostic tests for the presence of cryptic (hid-
den or hard to find) parasites been developed, but also vaccines may be
discovered that can protect people from such destructive protozoan dis-
eases as malaria. Malaria has killed more humans than any other disease in
history, and it currently causes the death of more than one million people,
and lowers the quality of life for millions of others, each year.
All species are involved in complex interrelationships with other spe-
cies that live in or on their bodies, or with which they intimately interact
behaviorally or ecologically. Such interactions may play a minor role in the
life and well-being of one or both of the associates, or they may be necessary
for the mutual survival of both. In relatively few symbiotic relationships,
one or both species may suffer damage or death. Pathogenic associations
are relatively rare, because disease or death of one symbiont generally re-
sults in corresponding disease or death of the other. Such relationships,
which cannot persist over evolutionarily long periods of time, may never-
theless cause catastrophic loss of life in nonadapted host populations.
All species are involved in complex interrelationships with other species
that live in or on their bodies, or with which they intimately interact
626
Symbiosis
behaviorally or ecologically. Such interactions may play a minor role in

the life and well-being of one or both of the associates, or they may be
necessary for the mutual survival of both. In relatively few symbiotic rela-
tionships, one or both species may suffer damage or death. Pathogenic as-
sociations are relatively rare, because disease or death of one symbiont
generally results in corresponding disease or death of the other. Such rela-
tionships, which cannot persist over evolutionarily long periods of time,
may nevertheless cause catastrophic loss of life in nonadapted host popu-
lations.
Domestic animals cannot live in some parts of the world, such as the
central portion of Africa, because they have little or no resistance to para-
sites of wild species, which are the normal hosts and are not harmed. Na-
tive species have coadapted with the parasites. This situation presents a
moral dilemma to humans. In the face of human needs for space and other
resources, should native animals be displaced or killed? Or should human
populations proactively slow their reproductive rates? History shows that
humanity has often chosen to take the former course.
The common view that animals which live in other animals are degen-
erate creatures that take advantage of more deserving forms of life is un-
derstandable but inaccurate. Symbionts are highly specialized animals
that do not live cost-free, or always to the detriment of their hosts. Symbi-
otic relationships between species have vastly increased the diversity,
complexity, and beauty of the living world.
Sneed B. Collard
See also: Animal-plant interactions; Biological invasions; Coevolution;

Communities: ecosystem interactions; Communities: structure; Competi-
tion; Ecology: definition; Food chains and webs; Lichens; Mycorrhizae;
Pollination; Predation; Trophic levels and ecological niches.

Boothroyd, John C., and Richard Komuniecki, eds. Molecular Approaches to
Parasitology. New York: Wiley-Liss, 1995.
Caullery, Maurice. Parasitism and Symbiosis. London: Sidgwick and Jack-
son, 1952.
Limbaugh, Conrad. “Cleaning Symbiosis.” Scientific American 205 (Au-
gust, 1961): 42-49.
Margulis, Lynn. “Symbiosis and Evolution.” Scientific American 225 (Au-
gust, 1971): 48-57.
Margulis, Lynn, and Dorion Sagan. Slanted Truths: Essays On Gaia, Symbio-
sis, and Evolution. New York: Copernicus, 1997.
627
Symbiosis
Noble, Elmer, Glenn Noble, Gerhard Schad, and Austin MacGinnes. Para-
sitology: The Biology of Animal Parasites. 6th ed. Philadelphia: Lea &
Febiger, 1989.
Toft, Catherine Ann, Andre Aeschlimann, and Liana Bolis, eds. Parasite-
Host Associations: Coexistence or Conflict? New York: Oxford University
Press, 1991.
Whitefield, Philip. The Biology of Parasitism: An Introduction to the Study of
Associating Organisms. Baltimore: University Park Press, 1979.
Zann, Leon P. Living Together in the Sea. Neptune City, N.J.: T. F. H., 1980.
Zinsser, Hans. Rats, Lice, and History. Reprint. New York: Bantam Books,
2000.
628
TAIGA
“Taiga” derives from a Russian word for the forests of cone-bearing, needle-leaved,
generally evergreen trees of northern Eurasia and North America. “Coniferous
forest” and “boreal forest” are other names given to this biome. Some botanists in-
clude the temperate rain forests along the Pacific Coast of North America and the
coniferous forests in the western mountains in the taiga.
W hile the term “coniferous forest” can be applied to temperate rain

forest and coniferous forest biomes in the western mountains, the
terms “taiga” and “boreal forest” should be restricted to the northern for-
ests. “Taiga” is also sometimes used in a more restricted way, to mean a
subdivision of the boreal forest.
Components
The dominant plants in the taiga are cone-bearing, needle-leaved, ever-
green trees, such as pines, spruces, and firs. North American taiga is domi-
nated by two species of spruce: black spruce (Picea mariana) and white
spruce (Picea glauca). Jack pine (Pinus banksiana), balsam fir (Abies balsamea),
and eastern larch (Larix laricina, a deciduous conifer) are also important in
parts of the taiga. A few deciduous flowering trees are also important com-
ponents. Quaking aspen (Populus tremuloides, the most widespread tree
species in North America) and paper birch (Betula papyrifera) are two ex-
amples. Eurasian taiga is dominated by related species of spruce and pine
and has the same character.
Determinants and Adaptations

Taiga occurs in a broad band across Canada, Alaska, Siberia, and Europe;
essentially, this band is interrupted only by oceans. This pattern suggests
that climate plays a major role in determining the distribution of the taiga.
Average temperatures are cool, and precipitation is intermediate, but
evaporation is low because of the cool temperatures. Hence, moisture is
generally available to plants during the growing season. The growing sea-
son is short, and winters are long. Permafrost is present in the northern
part of the taiga, and wetlands are common because drainage is often defi-
cient. These physical conditions are primarily determined by the high lati-
tude at which taiga occurs, but why taiga develops under these conditions
is not entirely clear.
629
Taiga
Taiga, or boreal forest, is

characterized by its location
in the northern latitudes
and its domination by cone-
bearing, evergreen trees,
notably spruces and pines.
A few deciduous, flowering
trees are also important
components, such as aspens,
the most widespread tree
species in the North
American taiga.
(PhotoDisc)
The length of the growing season may help explain why the dominant
taiga trees are evergreen. Because they retain their leaves through the win-
ter, these trees can carry out some photosynthesis on mild winter days.
More important, they avoid the energetic expense of replacing all their
leaves at one time. Deciduous trees put tremendous amounts of energy
into leaf replacement each spring and must replace those energy stores as
well as produce energy for growth during the growing season. Deciduous
forests generally occur south of the taiga, where the growing season is lon-
ger. However, some deciduous trees are successful in the taiga, so other ad-
aptations must also be important.
Asexual reproduction probably contributes to the success of taiga trees,
especially in severe environments. Black and white spruce reproduce by
layering, the growth of a new tree from a lower branch which makes con-
tact with the ground. Most deciduous trees of the taiga can sprout from the
roots or other underground parts if the aboveground part of the tree is
damaged or killed. Both strategies allow new trees to develop using the re-
sources of the parent tree. In contrast, some plants growing from seed do
not have sufficient resources to survive.
630
Taiga
Fire is an important environmental factor in the taiga. Many of the coni-

fers produce at least some cones which open and release their seeds only
after they have been heated intensely, as in a forest fire. Jack pine responds
to fire this way, as does black spruce to a lesser extent. Most deciduous
trees send up new stems from undamaged underground parts after a taiga
fire. White spruce does not employ either of these strategies but does have
efficient seed dispersal and so can move into a burned area fairly quickly.
Similar adaptations make Eurasian taiga species fit for life in northern en-
vironments. Apparently, no single suite of adaptations suits a tree species
for taiga life; instead various combinations of characteristics are employed
by the different species.
Adjacent Zones
The taiga is bordered by tundra to the north, and the meeting place be-
tween the two biomes is a broad transition zone often called the “taiga-
tundra,” or forest-tundra. This ecotone is composed of a mixture of forest
and tundra plants, with trees becoming fewer and smaller from south to
north until conditions become so harsh that trees can no longer grow.
The southern boundary of the taiga is often adjacent to deciduous for-
est, grassland, or parkland. These are also broad, transitional ecotones. In
eastern North America, the northern hardwood forest region is such a
transition zone and is composed primarily of a mixture of trees from the
deciduous forests and the taiga. The aspen parklands in the west are also
transitional. Quaking aspen from the taiga and grasses from western grass-
lands mix in this zone between the taiga and grassland biomes.
Environmental Concerns
Human activities may have less impact on the taiga than on many other
biomes, primarily because the taiga occurs in a harsh environment less ac-
cessible to humans than many other biomes. Still, there are serious con-
cerns. Acid rain became a problem for the taiga in eastern Canada in the
late twentieth century. These forests are northeast of the industrial centers
in the United States, and the prevailing southwesterly winds move nitro-
gen and sulfur oxides into eastern Canada, where they precipitate on
plants and soil. Both oxides interact with water to produce acids, thus acid-
ifying the soil and plant leaves. Many ecologists believe that acid precipita-
tion has seriously damaged the taiga of both North America and Eurasia.
Global warming is a second and perhaps more insidious threat to the
taiga. The taiga will almost certainly be negatively impacted by changes in
temperature, the length of the growing season, fire frequency and inten-
sity, and precipitation patterns. Taiga itself may play a role in carbon stor-
631
Taiga
age and mitigation of the greenhouse effect. This possibility, its role as a
source of timber, and the inherent value of the biome and its component
species make it imperative that the taiga be conserved.
Carl W. Hoagstrom

forests; Rain forests; Rain forests and the atmosphere; Rangeland; Reefs;
Savannas and deciduous tropical forests; Tundra and high-altitude biomes;
Wetlands.

Barbour, Michael G., and William Dwight Billings, eds. North American Ter-
restrial Vegetation. 2d ed. New York: Cambridge University Press, 2000.
Larsen, James A. The Boreal Ecosystem. New York: Academic Press, 1980.
Vankat, John L. The Natural Vegetation of North America: An Introduction.
Malabar, Fla.: Krieger, 1992.
632
TERRITORIALITY AND
AGGRESSION
Aggressive behavior and territoriality are common features of animals. Territories

may differ in function across species, but general trends occur. Territoriality is best
viewed as a means by which individuals maximize their own reproductive success
rather than as a mechanism of population regulation.
A ny field or forest inhabited by animals contains countless invisible

lines that demarcate territories of individuals of many different spe-
cies. Humans are oblivious to these boundaries yet have quick perception
of human property lines; other animals are equally oblivious to human de-
marcations. Most organisms, in fact, appear to attend only to the territorial
claims made by members of their own species. If separate maps of individ-
ual territories could be obtained for each species in the same habitat and
superimposed on one another, the resulting hodgepodge of boundaries
would show little consensus on the value of particular areas. Yet basic sim-
ilarities exist in why and how different species are territorial.
Causes of Territoriality
The existence of aggression and territorial behavior in nature hardly comes
as a surprise. Even casual observations at a backyard bird feeder reveal
that species that are commonly perceived as friendly can be highly aggres-
sive. The observation of birds at feeders can lead to interesting questions
concerning territorial behavior. For example, bird feeders usually contain
much more food than any one bird could eat: Why, then, are aggressive in-
teractions so common? Moreover, individuals attack conspecifics more of-
ten than birds of other species, even when all are eating the same type of
seeds.
Aggressive defense of superabundant resources is not expected to occur
in nature; however, bird feeders are not a natural phenomenon. Perhaps
the aggressive encounters that can be observed are merely artifacts of birds
trying to forage in a crowded, novel situation, or perhaps bird feeders in-
tensify aggressive interactions that occur less frequently and less conspicu-
ously in nature. While the degree to which aggression observed at feeders
mirrors reality is open to question, the observation of a greater intensity of
interactions between conspecifics definitely reflects a natural phenome-
633
Territoriality and aggression
non. Members of the same species are usually more serious competitors
than are members of different species because they exploit exactly the
same resources; members of different species might only share a few types
of resources. Despite the ecological novelty of artificial feeders, noting
which individuals win and lose in such an encounter can provide valuable
information on the resource-holding potential of individuals that differ in
various physical attributes such as body size, bill size, or even sex. For or-
ganisms that live in dense or remote habitats, this type of information can
often be obtained only by observations at artificial feeding stations.
Territorial defense can be accomplished by visual and vocal displays,
chemical signals, or physical encounters. The sequence of behaviors that an
individual uses is usually predictable. The first line of defense may involve
vocal advertisement of territory ownership. One function of birdsong is to
inform potential rivals that certain areas in the habitat are taken. If song
threats do not deter competitors, visual displays may be employed. If vi-
sual displays are also ineffective, then residents may chase intruders and, if
necessary, attack them. This sequence of behaviors is common in territorial
interactions because vocal and visual displays are energetically cheaper
than fighting and involve less risk of injury to the territory owner.
It may be less obvious why fighting is a necessary component in territo-
rial interactions for both territory owners and intruders. Without the threat
of bodily injury, there is no cost to intruders that steal the resources of an-
Two elephants exhibit territorial aggression. (Digital Stock)
634
other individual. This would severely hamper an owner’s ability to control

an area. On the other hand, if intruders never physically challenge territory
owners, then it would pay for all territory owners to exaggerate their abil-
ity to defend a resource. Thus, physical aggression may be essential. Ani-
mals do not frequently kill their opponents, however, so there must be
something that limits violence. Various species of animals possess formi-
dable weapons, such as large canine teeth or antlers, that are quite capable
of inflicting mortal wounds. Furthermore, a dead opponent will never
challenge again. Yet fights to the death are rare in nature. When they do oc-
cur, some novel circumstance is usually involved, such as a barrier that
prohibits escape of the losing individual. Restraint in normal use of weap-
ons, however, probably does not indicate compassion among combatants.
Fights to the death may simply be too costly, because they would increase
the chance that a victor would suffer some injury from a loser’s last desper-
ate attempts to survive.
Functions of Territory
Territories can serve various functions, depending on the species. For some,
the area defended is only a site where males display for mates; for others, it
is a place where parents build a nest and raise their offspring; for others, it
may be an all-purpose area where an owner can have exclusive access to
food, nesting sites, shelter from the elements, and refuge from predators.
These different territorial functions affect the area’s size and the length of
time an area is defended. Territories used as display sites may be only a
few meters across, even for large mammal species. Territorial nest sites
may be smaller still, such as the densely packed nest sites guarded by par-
ents of many colonial seabirds. All-purpose territories are typically large
relative to the body size of the organism. For example, some passerine
birds defend areas that may be several hundred meters across. Although
all three types of territories may be as ephemeral as the breeding season, it
is not uncommon for all-purpose territories to be defended year around.
The abundance and spatial distribution of needed resources determine
the economic feasibility of territoriality. On one extreme, if all required re-
sources are present in excess throughout the habitat, territory holders
should not have a reproductive advantage over nonterritory holders. At
the other extreme, if critical resources are so rare that enormous areas
would have to be defended, territory holders might again have no repro-
ductive advantage over nonterritory holders. If needed resources, how-
ever, are neither superabundant nor extremely rare and are somewhat
clumped in the habitat, territoriality might pay off. That is, territorial indi-
viduals might produce more offspring than nonterritorial individuals.
635
Studies of territoriality raise more questions than biologists can answer.

Researchers investigate how large an area an individual defends and
whether both sexes are equally territorial. They seek to determine whether
the territories of different individuals vary according to quality. The den-
sity of conspecifics may influence territoriality; on the other hand, terri-
toriality itself may serve to regulate population size, although evidence
suggests that this is an incidental effect.
All-purpose territories vary considerably in size, depending on the re-
source requirements of the individuals involved and the pattern of tem-
poral variation in resource abundance. In some organisms, individuals
only defend enough area to supply their “minimum daily requirements.”
In others, individuals defend a somewhat larger area—one that could still
support them even when resource levels drop. In others, individuals de-
fend territories that vary in size depending on current resource levels.
For example, pied wagtails (European songbirds) defend linear territo-
ries along riverbanks that are about six hundred meters long during the
winter. The emerging aquatic insects they consume are a renewable re-
source, but renewal rates vary considerably during the season. Rather
than adjusting territory size to match the current levels of prey abun-
dance in the habitat, wagtails maintain constant territory boundaries. This
inflexibility persists even though territories that extend for only three
hundred meters could adequately support an individual for about one-
third of the season. In contrast, the territory size of an Australian honey
eater varies widely during the winter. Nectar productivity of the flowers
visited by honey eaters varies considerably during the season. By adjust-
ing territory size to match changing resource levels, individual birds ob-
tain a relatively constant amount of energy each day (about eighteen
kilocalories).
Sex Roles
In some species, only males are territorial. In other species, both sexes de-
fend territories, but males defend larger territories than females do. In
some mammals in which both sexes are territorial, males are aggressive
only to other males, and females are only aggressive to other females. In
these species, male territories are sufficiently large to encompass the terri-
tories of several females. Presumably, these males have increased sexual
access to the females within their territories. Perhaps the most curious ex-
ample of sex-specific territorial behavior is observed in a number of coral
reef fish, in which all individuals in the population are initially female and
not territorial. As the individuals grow older and larger, some develop into
males. Once male, they engage in territorial behavior.
636
Within a species, significant variation in territory quality exists among

individuals. Studies on numerous species have demonstrated a relation-
ship between territory quality and an individual’s resource-holding poten-
tial. For example, larger individuals tend to control prime locations more
often than smaller individuals. In addition, possession of higher-quality
territories often results in increased reproductive success. For some spe-
cies, this occurs because individuals with better territories obtain mates
sooner or obtain more mates than individuals with poorer territories. In
other species, possession of superior territories increases the survival
chances of the owner.
As the density of conspecifics increases, the ability of individuals to
control territories decreases. In some species, the territorial system may
break down completely, with all individuals scrambling for their share of
needed resources in a chaotic fashion. In other species, the territorial sys-
tem is replaced by a dominance hierarchy. All competitors may remain in
the area, but their access to resources is determined by their rank in the hi-
erarchy. For example, dominant male elephant seals can successfully de-
fend from other males areas containing between eighty and one hundred
females. Very dense clusters of females, however (two hundred or more),
attract too many males for one male to monopolize. When this happens,
one male—usually the largest male—dominates the rest and maintains
disproportionate access to females.
Territoriality undeniably has an adaptive function: to increase the sur-
vival and reproductive success of individuals. Territoriality can also have
several possible incidental effects, one of which was once considered to be
an adaptive function: serving as a means of population regulation. The rea-
soning behind this hypothesis is simple. The number of territories in a hab-
itat would limit the number of reproducing individuals in a population
and would thereby prevent overpopulation that could cause a population
crash. Support for this hypothesis would include demonstration that a sig-
nificant number of nonbreeding adults exist in a population. Indeed, for
several species, experimental removal of territory owners has revealed
that “surplus” individuals quickly fill the artificially created vacancies. In
most of the species studied, however, these surplus individuals are pri-
marily males. Population growth can be curbed only by limiting the num-
ber of breeding females, not the number of breeding males. Furthermore,
the population regulation argument assumes that some individuals ab-
stain from reproduction for the good of the population. If such a popula-
tion did exist, a mutant individual that never abstained from reproducing
would quickly spread, and its descendants would predominate in future
generations.
637
Territoriality in the Field

Territoriality is typically investigated in the field using an observational
approach. Initial information collected includes assessing the amount of
area used by each individual, how much of that area is defended from
conspecifics, and exactly what is being defended. It is relatively easy to dis-
cern the spatial utilization of animals. For many species, all that is required
is capturing each individual, marking it for field identification, and watch-
ing its movements. For species that range long distances, such as hawks or
large mammals, and species that are nocturnal, radio telemetry is fre-
quently used. This methodology requires putting radio transmitters on the
individuals to be followed and using hand-held antennas, or antennas at-
tached to cars or airplanes, to monitor movements. For fossorial species
(animals that are adapted for digging), animal movements are often deter-
mined by repeated trapping. This method involves placing numerous
baited live traps above the ground in a predetermined grid.
Knowing the spatial utilization of an animal does not document ter-
ritoriality. Many types of animals repeatedly use the same regions in the
habitat but do not defend these areas from conspecifics. Such “home
ranges” may or may not contain areas that are defended (that is, territo-
ries). Territorial defense can be readily documented for some animals by
simply observing individual interactions. These data often need to be
supplemented by experiments. Behavioral interactions might occur only
in part of the organism’s living space because neighbors do not surround
it. For these individuals, researchers play tape-recorded territorial vocal-
izations or place taxidermy mounts of conspecifics in different locations
and note the response of the territory holder. For other species, such as
fossorial rodents, direct estimates of territory size cannot be obtained
because aggressive interactions cannot be observed; as a result, terri-
tory boundaries must be inferred from trapping information. Regions in
which only the same individual is repeatedly trapped are likely to be areas
that the individual defends. This is an indirect method, however, and can
be likened to watching the shadow of an organism and guessing what it is
doing.
It is often difficult to determine exactly what an animal is defending in
an all-purpose territory where organisms use many different types of re-
sources. Which resource, that is, constitutes the “reason” for territorial de-
fense? On the other hand, several resources may contribute in some com-
plex way. For many species these things simply are not known. This
uncertainty also complicates estimates of territory quality. For example,
red-winged blackbirds in North America have been particularly well stud-
ied for several decades by different investigators in various parts of the
638
species range. Males defend areas in marshes (or sometimes fields), and
some males obtain significantly more mates than others. Biologists think
that males defend resources that are crucial for female reproduction. Some
males may be more successful at mating than others because of variation in
territory quality. Yet the large number of studies done on this species has
not yielded a consensus on what the important resources are, whether
food, nest sites, or something else.
Theoretical investigations of territorial behavior often employ opti-
mality theory and game theory approaches. Optimality theory considers
the benefits and costs of territorial defense for an individual. Benefits and
costs might be measured simply as the number of calories gained and lost,
respectively. Alternatively, benefits might be measured as the number of
young produced during any one season; costs might be measured as the re-
duction in number of future young attributable to current energy expendi-
tures and risks of injury. For territorial behavior to evolve by means of nat-
ural selection, the benefits of territorial behavior to the individual must
exceed its costs.
Game theory analyses compare the relative success of individuals using
alternative behaviors (or “strategies”). For example, two opposing strate-
gies might be “defend resources from intruders” and “steal resources as
they are encountered.” In the simplest case, if some individuals only de-
fend and other individuals only steal resources, the question would be
which type of individual would leave the most offspring. Yet defenders in-
teract with other defenders as well as with thieves, and the converse holds
for thieves. By considering the results of interactions within and between
these two types of individuals, a game theory analysis can predict the
conditions under which one strategy would “win” or “lose” and how the
success of each type of individual would vary as the frequency of the other
increases in the population. A complete understanding of territoriality in-
volves not only empirical approaches in the field but also the development
of testable theoretical models. Considerable advances have been made re-
cently merging these two methodologies. Future investigations will no
doubt include experimental control over resource levels that will allow de-
finitive tests of predictions of alternative theoretical models.
Evolution of Terrioriality
Among animals in general, some species are highly aggressive in defend-
ing their living space, and others ignore or tolerate conspecifics in a nearly
utopian manner. Some animals are territorial during only part of the an-
nual cycle, and some only in specific areas that they inhabit; others remain
aggressive at any time and in any place. Thus, a main goal for researchers is
639
to unravel the ecological and evolutionary conditions that favor aggressive

behavior and territoriality.
Aggression and territorial behavior appear to have evolved in various
organisms because, in the past, aggressive and territorial individuals out-
reproduced nonaggressive and nonterritorial ones.
An implicit assumption of behavioral biologists is that animals other
than humans do not interact aggressively because of conscious reasoning,
nor are they consciously aware of the long-term consequences of aggres-
sive acts. Should these consequences be detrimental, natural selection will
eliminate the individuals involved, even if this means total extinction of
the species. Humans are different. They are consciously aware of their ac-
tions and of the consequences of such actions. They need only use con-
scious reasoning and biological knowledge of aggressive behavior to cre-
ate conditions that can reduce conflict between individuals and groups.
Richard D. Howard

Ethology; Habituation and sensitization; Hierarchies; Insect societies;
Mammalian social systems; Mimicry; Pheromones; Predation; Reproduc-
tive strategies.

Alcock, John. Animal Behavior. 7th ed. Sunderland, Mass.: Sinauer Associ-
ates, 2001.
Allen, Colin, and Marc Bekoff. Species of Mind: The Philosophy and Biology of
Cognitive Ethology. Cambridge, Mass.: MIT Press, 1997.
Davies, Nicholas B., and John R. Krebs. An Introduction to Behavioral Ecol-
ogy. 4th ed. Boston, Mass.: Blackwell Scientific Publications, 1997.
Dennen, J. van der, and V. S. E. Falger, eds. Sociobiology and Conflict: Evolu-
tionary Perspectives on Competition, Cooperation, Violence, and Warfare.
New York: Chapman and Hall, 1990.
Howard, Eliot. Territory in Bird Life. New York: Atheneum, 1962.
Ratcliffe, Derek A. The Peregrine Falcon. 2d ed. San Diego, Calif.: Academic
Press, 1993.
Wilson, Edward O. Sociobiology. Cambridge, Mass.: The Belknap Press of
Harvard University Press, 1975.
640
TROPHIC LEVELS AND ECOLOGICAL
NICHES
Types of ecology: Community ecology; Ecoenergetics; Ecosystem
ecology
A trophic level is a position in the food pyramid occupied by an organism based on

its food relationships with other organisms: what it eats, and what eats it. An eco-
logical niche is the physical space in which an animal lives and all the interactions
with the other living organisms and components of its environment.
T he idea of the niche probably had its first roots in ecology in 1910. At
that time, Roswell Johnson wrote that different species utilize different
niches in the environment. He theorized that individuals of a particular
species are only in certain places because of food supply and environmen-
tal factors that limit their distribution in an area. Later, in 1924, Joseph
Grinnel developed his concept of niche that centered on an organism’s dis-
tribution having limits set on it by climatic and physical barriers. At the
same time, Charles Elton was defining his own idea of niche. His descrip-
tion of niche involved the way an organism makes its living—in particular,
how it gathers food.
The Food Pyramid

For many years, ecologists focused on Elton’s definition and referred to
niche in terms of an organism’s place in the food pyramid. The food pyra-
mid is a simplified scheme in which organisms interact with one another
while obtaining food. The food pyramid is represented as a triangle, of-
ten with four horizontal divisions, each division being a different trophic
level.
The base of the food pyramid is the first trophic level and contains the
primary producers: photosynthetic plants. At the second trophic level are
the primary consumers. These are the herbivores, such as deer and rabbits,
which feed directly on the primary producers. Secondary consumers are
found at the third trophic level. This third trophic level contains carni-
vores, such as the mountain lion. The members of the uppermost trophic
level are the scavengers and decomposers, including hyenas, buzzards,
fungi, and bacteria. The organisms in this trophic level break down all the
nutrients (such as carbon and nitrogen) in the bodies of plants and animals
and return them to the soil to be absorbed and used by plants.
641
Trophic levels and ecological niches
It should be noted that no ecosystem actually has a simple and well-

defined food pyramid. Many organisms interact with more than only the
organisms at the adjacent trophic levels. For example, a coyote could be
considered to belong to the third trophic level with the carnivores, but the
coyote also feeds on occasional fruits and other primary producers. Ba-
sically, all living things are dependent on the first trophic level, because it
alone has the capability to convert solar energy to energy found in, for ex-
ample, glucose and starch. The food pyramid takes the geometric form of a
triangle to show the flow of energy through a system.
Photosynthetic plants lose 10 percent of the energy they absorb from the
sun as they convert solar energy into glucose and starch. In turn, the herbi-
vores can convert and use only 90 percent of the energy they obtain by eat-
ing plants. Hence, less energy is found at each higher trophic level. Because
of this reduced energy, fewer organisms can be supported by each higher
trophic level. Consequently, the sections of the pyramid get smaller at each
higher trophic level, representing the decreasing levels of energy and num-
ber of members.
Types of Niches
Through the years, two concepts of niche have evolved in ecology. The first
is the place niche, the physical space in which an organism lives. The sec-
ond is the ecological niche, and it encompasses the particular location oc-
cupied by an organism and its functional role in the community.
The functional role of a species is not limited to its placement along a
food pyramid; it also includes the interactions of a species with other or-
ganisms while obtaining food. For example, the methods used to tolerate
the physical factors of its environment, such as climate, water, nutrients,
soils, and parasites, are all part of its functional role. In other words, the
ecological niche of an organism is its natural history: all the interactions
and interrelationships of the species with other organisms and the envi-
ronment.
The study of the interrelationships among organisms has been the focus
of ecological studies since the 1960’s. Before this time, researchers had fo-
cused on the food pyramid and its effect on population changes of merely a
single species. One example, the classic population study of the lynx and
the snowshoe hare of Canada, originally focused on the interactions of the
species in the food pyramid. It was discovered that the lynx had a ten-year
population cycle closely following the population cycle of its prey, the
snowshoe hare. The lynx population appeared to rise, causing a decline in
the population of the snowshoe hare. In the investigations that followed,
however, studies diverted the focus from the food pyramid to other ele-
642
The Food Pyramid:

Trophic Levels
Decomposers,
Scavengers:
Bacteria, Fungi,
Buzzards, Hyenas
Seconday Consumers:
Carnivores
Primary Consumers:
Herbivores
Primary Producers: Photosynthetic Plants
More Energy .................................................................. Less Energy
ments of the niche of the two species. For example, the reproductive nature
of the hare provided a contradiction to the simple predator-prey explana-
tion. The hare has a faster rate of reproduction than the lynx. It seemed im-
possible that the significantly lower population of lynx could effectively
place sufficient predator pressure on the hare to cause its drastic decline in
numbers. Therefore, it appeared that the population dynamics of the hare
and lynx were regulated by more than simply a predator-prey relation-
ship.
Later studies of the lynx and hare suggested that the peaks and dives in
the two populations may also be a factor of parasites of the hare that are
643
carried by the lynx. A rise in the lynx population increases the carriers of
parasites of the hare. Therefore, it is thought that, although the hare has a
much greater reproduction rate than the lynx, the population of hares will
still decline because of the combination of predation by the lynx and the in-
creased frequency of parasites of the hare. This study involved looking at
more than one dimension of the ecological niche of a species and broke
away from concentrating on only the interactions between organisms in
the food pyramid.
Niche Overlap: Interspecific

The goal of understanding how species interact with one another can also
be better accomplished by defining the degree of niche overlap, the degree
of the sharing of resources between two species. When two species use one
or more of the same elements of an ecological niche, they exhibit inter-
specific competition. It was once believed that interspecific competition
would always lead to survival of only the better competitor of the two spe-
cies. That was the original concept of the principle of the competition ex-
clusion law of ecology: No two species can utilize the same ecological
niche. It was conjectured that the weaker competitor would either migrate,
begin using another resource not used by the stronger competitor, or be-
come extinct. It is now believed that the end result of two species sharing
elements of ecological niches may not always be exclusion.
Ecologists theorize that similar species do, in fact, coexist, despite the
sharing of elements of their ecological niches, because of character dis-
placement, which leads to a decrease in niche overlap. Character displace-
ment involves a change in the morphological, behavioral, or physiological
state of a species without geographical isolation. Character displacement
occurs as a result of natural selection arising from competition between
one or more ecologically similar species. Examples might be changes in
mouth sizes so that they begin to feed on different sizes of the same food
type, thereby decreasing competition.
Specialists and Generalists

The more specialized a species, the more rigid it will be in terms of its eco-
logical niche. A species that is general in terms of its ecological niche needs
will be better able to find and use an alternative for the common element of
the niche. Since a highly specialized species cannot substitute whatever is
being used, it cannot compete as well as the other species. Therefore, a spe-
cialized species is more likely to become extinct.
For example, a panda is a very specialized feeder, eating mainly bam-
boo. If a pest is introduced into the environment that destroys bamboo, the
644
panda will probably starve, being unable to switch to another food source.
On the other hand, the coyote is a generalized feeder. A broad variety of
food types make up its diet. If humans initiate a pest-control program, kill-
ing the population of rabbits, the coyote will not fall victim to starvation,
because it can switch to feeding predominantly on rodents, insects, fruits,
and domesticated animals (including cats, dogs, and chickens). Hence,
species with specialized ecological niche demands (specialists) are more in
danger of extinction than those with generalized needs (generalists). Al-
though this fundamental difference in survival can be seen between spe-
cialists and generalists, it must be noted again that exclusion is not an inev-
itable result of competition. There are many cases of ecologically similar
species that coexist.
Niche Overlap: Intraspecific

When individuals of the same species compete for the same elements of
the ecological niche, it is referred to as intraspecific competition. Intra-
specific competition has the opposite effect of interspecific competition:
niche generalizations. In increasing populations, the first inhabitants will
have access to optimal resources. The opportunity for optimal resources
decreases as the population increases; hence, intraspecific competition in-
creases. Deviant individuals using marginal resources may slowly begin to
use less optimal resources that are in less demand. That can lead to an in-
crease in the diversity of ecological niches used by the species as a whole.
In other words, the species may become more generalized and exploit
wider varieties of niche elements.
Representing a situation on the opposite end of the spectrum from that
of two organisms competing for the same dimension of an ecological niche
is the vacant niche theory. This ecological principle states that when an or-
ganism is removed from its ecological niche, space, or any other dimension
of the niche, another organism of the same or similar species will reinvade.
Field Research
Theoretical studies of ecological niches are abstract, since humans are lim-
ited to three-dimensional diagrams, and there are more dimensions than
three to an ecological niche. This multidimensionality is referred to as the
n-dimensional niche. This abstract n-dimensional niche can be studied
mathematically and statistically, but the study of ecological niches is
mainly a field science. Therefore, its techniques are mainly those used for
field research.
Research that attempts to describe all the elements of the n-dimensional
ecological niche would require extensive observations. Yet, ecological
645
niches are difficult to measure not only because of the plethora of data that
would have to be collected but also because of the element of change in na-
ture. The internal and external environment of an organism is always dy-
namic. Nothing in life is static, even if equilibrium is established.
These constant fluctuations create daily and seasonal changes in space
and ecological niches. Therefore, because of the constant fluctuations,
merely descriptive field observations would not be reliable depictions of
an organism’s ecological niche. Ecologists must also resort to quantitative
data of measurable features of an organism’s ecological niche. For exam-
ple, the temperature, pH, light intensity, algae makeup, predators, and
activity level of the organism are measurable features of an ecological
niche in a pond community. The difficulty is in the collection of each of
the necessary measurements making up an ecological niche. The ecolo-
gist would have to limit the data to a manageable number of specific di-
mensions of the niche based on conjecture and basic intuition. Such limi-
tations often lead to incomplete and disconnected measurements that can
at best only partially describe a few of the dimensions of the ecological
niche.
Ecologists realize that complete observations and measurements of all
the dimensions of an organism’s ecological niche are unattainable. The
focus in understanding how a species interacts with its community cen-
ters on determining the degree of niche overlap between any two species.
In other words, the level of competition for space niche and resources.
Studies of this niche overlap are typically limited to dimensions that can
be quantitatively measured. Yet, there is still the problem of deciding
which of the dimensions are involved in the competition between the two
species. Again, the ecologist must usually rely on inherent knowledge
about the two species in question. Often, researchers investigating niche
competition measure no more than four ecological niche dimensions to de-
termine the niche overlap in an attempt to understand how two individu-
als competing for the same space, resources, or other ecological niche fea-
tures can coexist.
Field methods for observations and quantitative measurements of ele-
ments of ecological niches, niche overlap, and niche competition are prob-
ably endless. To name a few, describing an organism’s niche may involve
fecal samples to determine its diet, fecal samples of possible predators to
identify its primary predator, animal and plant species checklists of its
space niche along with soil components, climatic trends, and the like.
Niche competition and overlap often can be studied first in the laboratory
under controlled situations. One method might involve recording the pop-
ulation dynamics of the species as different elements in the ecological
646
niche are manipulated to determine which is the better competitor and

what is the resource that is most responsible for limiting the population
size.
Niche and Community

The shift in meaning and study from merely space and trophic level place-
ment in the food pyramid to ecological niche of n dimensions has been
beneficial for the field of ecology. This focus on community ecology is obvi-
ously much more productive for the goal of ecology, the understanding of
how all living organisms interact with one another and with nonliving ele-
ments in the environment.
Perhaps more important is the attempt to describe niches in terms of
community ecology, which can be essential for some of humankind’s con-
frontations with nature. For example, it has become increasingly apparent
that synthetic chemicals are often too costly and too hazardous to continue
using for control of crop pests and carriers of diseases. The goal is to con-
trol pests effectively with biological controls. Biological controls can in-
volve the introduction of natural predators of the undesirable pest or the
introduction of a virus or bacteria that eliminates the pest and is harmless
to humans and wildlife.
The success of a biological control is directly proportional to the knowl-
edge of the pest’s n-dimensional ecological niche and the other organisms
with which it comes in contact. A classic example of the havoc that can re-
sult from manipulations of nature without adequate ecological informa-
tion is when Hawaii attempted to use biological controls to eradicate a
population of snakes, which humans had accidentally introduced. The bio-
logical control used was the snake’s natural predator, the mongoose. One
very important dimension of the ecological niche of both species was ig-
nored. One species was active only at night, while the other was active only
during the day. Needless to say, this particular venture with a biological
control was not a success.
Another relevant function of community-oriented studies of ecological
niches involves endangered species. In addition to having aesthetic and
potential medicinal values, an endangered organism may be a keystone
species, a species on which the entire community depends. A keystone
species is so integral to keeping a community healthy and functioning that
if obliterated the community no longer operates properly and is not pro-
ductive.
Habitat destruction has become the most common cause of drastic pop-
ulation declines of endangered species. To enhance the habitat of the en-
dangered species, it is undeniably beneficial to know what attracts a spe-
647
cies to its particular preferred habitat. This knowledge involves the details
of many of the dimensions of its ecological niche integral to its population
distribution. Another common means of endangering the survival of a
species is to introduce an organism or exotic species that competes for the
same resources and displaces the native species. Solving such competition
between native and introduced species would first involve determining
niche overlap.
It is often stated that an ounce of prevention is worth a pound of cure.
Thus, the researching and understanding of all the dimensions of ecologi-
cal niches are key to preventing environmental manipulations by human-
kind that might lead to species extinction. Many science authorities have
agreed that future research in ecology and related fields should focus on
solving three main problems: species endangerment, soil erosion, and
solid waste management.
This focus on research in ecology often means that studies of pristine
communities, those undisturbed, will be the most helpful for future resto-
ration projects. Although quantitative and qualitative descriptions of pris-
tine areas seem to be unscientific at the time they are made, because there is
no control or experimental group, they are often the most helpful for later
investigations. For example, after a species has shown a drastic decline in
its population, the information from the observations of the once-pristine
area may help to uncover what niche dimension was altered, causing the
significant population decrease.
Jessica O. Ellison
See also: Animal-plant interactions; Balance of nature; Biodiversity; Bioge-

ography; Biological invasions; Coevolution; Communities: ecosystem in-
teractions; Communities: structure; Competition; Food chains and webs;
Herbivores; Omnivores; Phytoplankton; Predation; Symbiosis.

Bronmark, Christopher, and Lars-Anders Hansson. The Biology of Lakes and
Ponds. New York: Oxford University Press, 1998.
Ehrlich, Paul R. “Who Lives Together, and How.” In The Machinery of Na-
ture. New York: Simon & Schuster, 1986.
Giller, Paul S. Community Structure and the Niche. New York: Chapman and
Hall, 1984.
Odling-Smee, F. John, Kevin N. Laland, and Marcus W. Feldman. “Niche
Construction.” American Naturalist 147, no. 4 (April, 1996): 641-649.
Rayner, Alan D. M. Degrees of Freedom: Living in Dynamic Boundaries. River
Edge, N.J.: World Scientific Publications, 1997.
648
Ricklefs, Robert E. Ecology. 4th ed. New York: Chiron Press, 1999.
Shugart, Herman H. Terrestrial Ecosystems in Changing Environments. New
Smith, Robert L. Ecology and Field Biology. 6th ed. San Francisco: Benjamin/
Cummings, 2001.
Stone, Richard. “Taking a New Look at Life Through a Functional Lens.”
Science 269, no. 5222 (July, 1995): 316-318.
649
TROPISMS
Type of ecology: Physiological ecology
Tropisms represent a variety of adaptations of plants that have allowed them to

grow toward or away from environmental stimuli such as light, gravity, objects to
climb, moisture in soil, or the position of the sun. These rapid responses, which
make use of separate systems for detecting and responding to stimuli, help a plant
to survive in its particular habitat.
A lthough plants appear not to move, they have evolved adaptations to

allow movement in response to various environmental stimuli; such
mechanisms are called tropisms. There are several kinds of tropism, each
of which is named for the stimulus that causes the response. For exam-
ple, gravitropism is a growth response to gravity, and phototropism is a
growth response to unidirectional light. Tropisms are caused by differen-
tial growth meaning that one side of the responding organ grows faster
than the other side of the organ. This differential growth curves the organ
toward or away from the stimulus. Growth of an organ toward an environ-
mental stimulus is called a positive tropism; for example, stems growing
toward light are positively phototropic. Conversely, curvature of an organ
away from a stimulus is called a negative tropism. Roots, which usually
grow away from light, are negatively phototropic. Tropisms begin within
thirty minutes after a plant is exposed to the stimulus and are usually com-
pleted within approximately five hours.
Phototropism
Phototropism is a growth response of plants to light coming from one di-
rection. Positive phototropism of stems results from cells on the shaded
side of a stem growing faster than cells along the illuminated side; as a re-
sult, the stem curves toward the light. The rapid elongation of cells along
the shaded side of a stem is controlled by a plant hormone called auxin that
is synthesized at the stem’s apex. Unidirectional light causes the auxin to
move to the shaded side of stems. The increased amount of auxin on the
shaded side of stems causes cells there to elongate more rapidly than cells
on the lighted side of the stem. This, in turn, causes curvature toward the
light.
Only blue light having a wavelength of less than 500 nanometers can in-
duce phototropism. The photoreceptors in this system are called crypro-
chromes and may alter the transport of auxin across cellular membranes,
650
Tropisms
thereby facilitating its transport to the shaded side of the stem. Photo-
tropism is important for two main reasons: It increases the probability of
stems and leaves intercepting light for photosynthesis and of roots obtain-
ing water and dissolved minerals that they need.
Gravitropism
Gravitropism is a growth response to gravity. The positive gravitropism of
roots involves the root cap, a tiny, thimble-shaped organ approximately 0.5
millimeter long that covers the tip of roots. Decapped roots grow but do
not respond to gravity, indicating that the root cap is necessary for root
gravitropism. Gravity-perceiving cells, called columella cells, are located
in the center of the root cap. Each columella cell contains fifteen to twenty-
five amyloplasts (starch-filled plastids) which, under the influence of grav-
ity, sediment to the lower side of columella cells. This gravity-dependent
sedimentation of amyloplasts is the means whereby roots sense gravity,
possibly by generating electrical currents across the root tip. These gravity-
induced changes are then transmitted to the root’s elongating zone, lo-
Tropisms
Gravitropism (geotropism) Phototropism
Heliotropism (solar tracking) Thigmotropism
651
Tropisms
cated 3 to 6 millimeters behind the root cap. The differential growth that
causes curvature occurs in the elongating zone.
When roots are oriented horizontally, growth along the lower side of the
elongating zone is inhibited, thereby causing the root to curve downward.
Among the first events that produce this differential growth is the accumu-
lation of calcium ions along the lower side of the root tip. Calcium ions
move to the lower side of the cap and elongating zone of horizontally ori-
ented roots. This movement may be aided by electrical currents in the root.
The accumulation of calcium along the lower side of the root causes the
auxin to accumulate there as well. Because auxin inhibits cellular elonga-
tion in roots, the lower side of the root grows slower than the upper side of
the root, and the root curves downward. When the root becomes vertical,
the lateral asymmetries of calcium and auxin disappear, and the root
grows straight down.
Gravity-sensing cells in stems are located throughout the length of the
stem. As in roots, the auxin and calcium ions in stem cells direct the nega-
tive gravitropism (in this case, upward curvature) of shoots. As auxin ac-
cumulates along the lower side, calcium ions gather along the upper side
of horizontally oriented stems. The accumulation of auxin along the stem’s
lower side stimulates cellular elongation there. Gravitropism increases the
probability of two important results: Roots will be more likely to encounter
water and minerals, and stems and leaves will be better able to intercept
light for photosynthesis.
Thigmotropism
Thigmotropism is a growth response of plants to touch. The most common
example of thigmotropism is the coiling exhibited by specialized organs
called tendrils. Tendrils are common on twining plants such as morning
glory and bindweed. Prior to touching an object, tendrils often grow in a
spiral. This type of growth is called circumnutation, and it increases the ten-
dril’s chances of touching an object to which it can cling. Contact with an
object is perceived by specialized epidermal cells on the tendril. When the
tendril touches an object, these epidermal cells control the differential
growth of the tendril. This differential growth can result in the tendril com-
pletely circling the object within five to ten minutes. Thigmotropism is often
long-lasting. For example, stroking one side of a tendril of garden pea for
only a few minutes can induce a curling response that lasts for several days.
Thigmotropism is probably controlled by auxins and ethylene, as these reg-
ulate thigmotropic-like curvature of tendrils even in the absence of touch.
Growing tendrils touched in the dark do not respond until they are illu-
minated. This light-induced expression of thigmotropism may indicate a
652
Tropisms
requirement for adenosine triphosphate (ATP), as ATP will substitute for

light in inducing thigmotropism of dark-stimulated tendrils. Tendrils can
store the sensory information received in the dark, but light is required for
the coiling growth response to occur. Thigmotropism by tendrils allows
plants to “climb” objects and thereby increases their chances of intercept-
ing light for photosynthesis.
Hydrotropism and Heliotropism

Roots also grow toward wet areas of soil. Growth of roots toward soil
moisture is called hydrotropism. Roots whose caps have been removed do
not grow toward wet soil, suggesting that the root cap is the site of mois-
ture perception by roots. Hydrotropism is probably controlled by interac-
tions of calcium ions and hormones such as the auxins.
Heliotropism, or “solar tracking,” is the process by which plants’ or-
gans track the relative position of the sun across the sky, much like a ra-
dio telescope tracks stars or satellites. Different plants have different types
of heliotropism. The “compass” plants (Lactuca serriola and Silphium
laciniatum) that grow in deserts orient their leaves parallel to the sun’s rays,
thereby decreasing leaf temperature and minimizing desiccation. Plants
that grow in wetter regions often orient their leaves perpendicular to the
sun’s rays, thereby increasing the amount of light intercepted by the leaf.
Heliotropism occurs in many plants, including cotton, alfalfa, and beans.
Sunflowers get their name from the fact that their flowers follow the sun
across the sky. On cloudy days, leaves of many heliotropic plants become
oriented horizontally in a resting position. If the sun appears from behind
the clouds late in the day, leaves rapidly reorient themselves—they can
move up to 60 degrees in an hour, which is four times more rapid than the
movement of the sun across the sky. Heliotropism is controlled by many
factors, including auxins.
Practical Implications
Biologists are studying tropisms in hopes of being able to mimic these de-
tection and “guidance” systems for human use. Scientists at the National
Aeronautics and Space Administration (NASA), for example, have been
studying the way plants perceive and respond to gravity to learn how to
grow plants in deep space. Understanding the gravity detection and guid-
ance systems in plants may help people design more effective rockets,
which, like plants, must respond to gravity to be effective.
Randy Moore
See also: Adaptations and their mechanisms.
653
Tropisms

Campbell, Neil A., and Jane B. Reece. Biology. 6th ed. San Francisco:
Benjamin Cummings, 2002.
Evans, Michael L., Randy Moore, and Karl H. Hasenstein. “How Roots Re-
spond to Gravity.” Scientific American 255 (December, 1986): 112-119.
Hart, James Watnell. Plant Tropisms and Other Growth Movements. Boston:
Unwin Hyman, 1990.
Haupt, W., and M. E. Feinleib, eds. The Physiology of Movements. New York:
Salisbury, Frank B., and Cleon W. Ross. Plant Physiology. 4th ed. Belmont,
Calif.: Wadsworth, 1992.
Satter, R. L., and A. W. Galston. “Mechanisms of Control of Leaf Move-
ments.” Annual Review of Plant Physiology 32 (1981): 83-103.
Taiz, Lincoln, and Eduardo Zeiger. Plant Physiology. 2d ed. Sunderland,
Mass.: Sinauer Associates, 1998.
654
TUNDRA AND HIGH-ALTITUDE
BIOMES
Regions where no trees grow because of frozen soil or extreme water runoff due to
steep grades (at high altitudes) are known as tundra. High altitude biomes have
similar limitations on the growth of plant life.
T undra landscapes appear where long, cold winters, a permanently

frozen subsoil, and strong winds combine to prevent the development
of trees. The resulting landscapes tend to be vast plains with low-growing
forbs and stunted shrubs. Vast areas of this biome encircle the northern-
most portions of North America and Eurasia, constituting the Arctic tun-
dra. Climatic conditions atop high mountains at all latitudes are similar;
these small, isolated areas are called the alpine tundra.
Permafrost
The low temperatures of the tundra regions cause the formation of a per-
manently frozen layer of soil known as permafrost. Characteristic of Arctic
tundra, permafrost, which varies in depth according to latitude, thaws at
the surface during the brief summers. As the permafrost below is impene-
trable by both water and plant roots, it is a major factor in determining the
basic nature of tundra.
The alternate freezing and thawing of soil above the permafrost creates
a symmetrical patterning of the land surface characteristic of Arctic tun-
dra. Perhaps the best known features of the landscape are stone polygons
that result when frost pushes larger rocks toward the periphery, with
smaller ones occupying the center of each unit. This alteration of the tun-
dra landscape, called cryoplanation, is the major force in molding Arctic
tundra landscapes.
In contrast, alpine tundra generally has little or no permafrost. Even
though alpine precipitation is almost always higher than for Arctic tundra,
steep grades result in a rapid runoff of water. Alpine soils are, therefore,
much drier, except in the flat alpine meadows and bogs, where conditions
are more like those of Arctic areas.
Vegetation
Both Arctic and alpine tundra regions are composed of plants that have
adapted to the same generally stressful conditions. Biodiversity of both
655
Tundra and high-altitude biomes
plants and animals—the total number of species present—is low com-

pared to most other ecosystems. Plant growth is slow because of the short
growing seasons and the influence of permafrost. Most tundra plants are
low-growing perennials that reproduce vegetatively rather than by seed.
Often they grow in the crevices of rocks that both shelter them in the winter
and reflect heat onto them in summer.
Common plants of the low-lying Arctic tundra sites include various
sedges, especially cottongrass, and sphagnum moss. On better-drained
sites, biodiversity is higher, and various mosses, lichens, sedges, rush spe-
cies, and herbs grow among dwarfed heath shrubs and willow. The ar-
rangement of plants within a small area reflects the numerous microcli-
mates resulting from the peculiar surface features.
Alpine plants possess many of the features of Arctic plants. However,
because strong winds are such a prominent feature of the alpine environ-
ment, most of the plants grow flat on the ground, forming mats or cush-
ions.
Below alpine tundra and south of Arctic tundra, there is the boreal (also
known as taiga) biome, dominated by coniferous forest. Between the forest
and tundra lies a transitional zone, or ecotone. This ecotone is character-
ized by trees existing at their northern (or upper) limit. Especially in alpine
regions, stunted, gnarled trees occupy an area called krummholz. In North
Image Not Available
656
America, the krummholz is much more prominent in the Appalachian

Mountains of New England than in the western mountains.
Conservation
Like all world biomes, tundra regions are subject to degradation and de-
struction, especially as a result of human activities. Because of low human
population density and their unsuitability for agriculture, tundras gener-
ally are less impacted by humans than are grasslands and forests. How-
ever, tundra ecosystems, when disturbed, recover slowly, if at all. As most
tundra plants lack the ability to invade and colonize bare ground, the pro-
cess of ecological succession that follows disturbances may take centuries.
Even tire tracks left by vehicles can endure for decades. The melting of per-
mafrost also has long-lasting effects.
The discovery of oil and gas in tundra regions, such as those of Alaska
and Siberia, has greatly increased the potential for disturbances. Heavy
equipment used to prospect for fossil fuels and to build roads and pipe-
lines has caused great destruction of tundra ecosystems. As the grasses
and mosses are removed, the permafrost beneath melts, resulting in soil
erosion. The disposal of sewage, solid wastes, and toxic chemicals poses
special problems, as such pollutants tend to persist in the tundra environ-
ment longer than in warmer areas.
Animals of the Arctic tundra, such as caribou, have been hunted by
the native Inuit using traditional methods for centuries without an im-
pact on populations. The introduction of such modern inventions as snow-
mobiles and rifles has caused a sharp decline in caribou numbers in some
areas.
Although efforts at restoring other ecosystems, especially grasslands,
have been quite successful, tundra restoration poses difficult problems.
Seeding of disturbed Arctic tundra sites with native grasses is only mar-
ginally successful, even with the use of fertilizers. In alpine tundra, restora-
tion efforts have been somewhat more successful but involve transplant-
ing as well as seeding and fertilizing. A recognition of natural successional
patterns and long-term monitoring is a necessity in such efforts.
Thomas E. Hemmerly
See also: Biomes: determinants; Biomes: types; Forests; Habitats and

biomes; Mountain ecosystems; Taiga.

Johnson, Rebecca L. A Walk in the Tundra. Minneapolis: Carolrhoda Books,
2000.
657
Sayre, April Pulley. Tundra. Frederick, Md.: Twenty-first Century Books,

1995.
Shepherd, Donna W. Tundra. New York: Franklin Watts, 1997.
Smith, R. L., and T. M. Smith. Elements of Ecology. 4th ed. San Francisco:
Benjamin Cummings, 1998.
Walker, Tom. Caribou: Wanderer of the Tundra. Portland, Oreg.: Graphic Arts
Center, 2000.
Zwinger, Ann H., and Beatrice E. Willard. Land Above the Trees: A Guide to
American Alpine Tundra. Boulder, Colo.: Johnson Books, 1996.
658
URBAN AND SUBURBAN WILDLIFE
Types of ecology: Landscape ecology; Restoration and conservation
ecology
The global increase in the human population growth and density has seen a simul-
taneous increase in the growth of urban and suburban areas throughout the world.
As urban habitats and their suburban extensions have become more common, the
wildlife of these human landscapes has become the focus of attention and study.
A nimals and plants of cities and suburbs are categorized as urban

wildlife. As cities and suburbs grow ever larger and displace natural
habitats, many city and suburban landscapes have become more attractive
for certain kinds of wildlife, or at least urban wildlife has become more no-
ticeable. Urban wildlife consists of an eclectic and unlikely mix of escaped
pets (including cats, dogs, reptiles, exotics, and caged birds), feral animals,
furtive and temporary intruders from adjacent natural habitats, and spe-
cies whose natural ecology and behavior enable them to fit within human-
modified landscapes and tolerate living in close proximity to humans.
Urban Habitats
Urban landscapes present a seemingly stark and forbidding environment
for wildlife. The horizontal streets and sidewalks are punctuated by rising
angles and arches of concrete and steel that in turn are topped by wood and
metal rooftops. Overhead, a maze of telephone, power, and cable lines lim-
its vertical movement, while vehicle and foot traffic poses a constant threat
to surface movement. All of these edifices and connecting corridors and
lines result in a complex, vertically structured environment that some ani-
mals find difficult to maneuver yet to which other animals quickly adapt.
In addition to this monotonous and often dangerous structural diver-
sity, urban wildlife is subject to elevated and often almost continuous noise
and disturbance and is constantly exposed to an enormous variety of resi-
dential wastes (garbage, litter, excess water, salts, sewage), vehicular pol-
lutants (lubricants, greases, gasoline, hydrocarbons, nitrogen oxides), and
chemical wastes (pesticides, paints, lead, mercury, contaminants).
Despite the forbidding features of urban habitats, a surprising variety of
wildlife manages to exist on a more or less permanent basis. In fact, some
kinds of wildlife can be found even in the midst of the most degraded forms
of urban blight. Ailanthus, which is also commonly called tree-of-heaven, is
but one of many opportunistic trees and shrubs that can take root and
659
Urban and suburban wildlife
grow given a bare minimum of soil and nutrients. A simple linear crack in
the pavement of a sidewalk, a little-used roadway, an unused parking area,
or a vacant lot can trap enough windswept dirt to offer a growing substrate
for Ailanthus and similar hardy plants. Each Ailanthus, in turn, provides
food and shelter for equally tough and adaptable wildlife, ranging from
the variety of invertebrates that colonize and feed upon Ailanthus to birds
and mammals that take shelter or find food in its branches and foliage.
Similarly, every invading sprig of grass, wildflower, shrub, or tree, how-
ever large or small, creates its own suite of microhabitats which, in turn, of-
fer colonization opportunities for other plants and animals, the whole ulti-
mately contributing to an overall increase in urban biodiversity.
Benefits of Urban Habitats

Ailanthus is an example of those plants and animals able to tolerate the most
extreme urban conditions, but in reality most urban wildlife derives a number
of benefits by living within the confines of cities and suburbs. Far from being
homogeneous expanses of concrete, most urban centers are a patchwork of
different habitats—residential, commercial, and industrial buildings, ware-
houses, power stations, vacant lots, detached gardens, rooftop gardens,
and alleyways—that each offer innumerable opportunities for wildlife.
Many urban areas also have a number of limited access areas that animals
are quick to adopt for shelter and breeding places; these include fenced-in
lots and boarded-up buildings, along with a rabbit warren of underground
tunnels, ducts, steam and water pipes, basements, and access ways.
City lights extend foraging time and opportunities, allowing wildlife to
hunt for food not only throughout the day but also during much of the
night, as needed. Urban nooks and crannies offer an extensive variety of
microhabitats that differ fundamentally in size, microclimate, and struc-
tural features. These microhabitats serve primarily as shelters and breed-
ing sites for city wildlife. Many birds, such as house sparrows (Passer
domesticus) and Eurasian starlings (Sturnis vulgaris) nest in innumerable
crevices, cracks, nooks, niches, and sheltered rooftops. Pigeons (Columbia
livia) and starlings hide in sheltered enclaves offered by bridge abutments
and supports, archways, and other edifices.
The most adaptable forms of wildlife are quick to find and take advan-
tage of subtle advantages offered in urban habitats: Many birds cluster
around chimneys and roof reflectors or in shelters afforded by lee sides of
rooftops during harsh cold and windstorms. Others are equally quick to
obtain warmth by sitting on poles, rooftops, or other elevated perches to
orient toward sunlight, while at ground level animals gather near gratings,
vents, and underground heating pipes.
660
Urban Scavengers
Urban wildlife just as quickly concentrates in areas where potential food
is made available—for instance, during trash pickup—then just as quickly
disperses to find new food sources. Most forms of urban wildlife for-
age opportunistically as scavengers, specializing in finding and consum-
ing all bits of discarded food, raiding trash cans, and concentrating at
waste collection and disposal centers. Thus, the rubbish dumps, found
in or immediately adjacent to every city of the world, attract an amaz-
ing diversity of small mammals and birds. Feeding on the scavenged
food of urban areas and bird feeders is much more efficient because it re-
quires less energy to find or catch and is usually available throughout the
year.
Because of the need to find and exploit temporary food resources, some
of the most successful urban animals forage in loose groupings or flocks:
The more eyes there are for searching, the more feeding opportunities can
be identified and exploited. Solitary and nonsocial species often do less
well in urban environments simply because they lack the collective power
of the group to find food and shelter, and avoid enemies.
The availability of a year-round food supply—however tenuous and
temporary—along with the presence of an enormous variety of safe shel-
ters and breeding sites promotes a higher life expectancy, which partly or
mostly balances the higher vehicle-related death rates to which urban
wildlife is continuously subject.
Urban Parks
Parks and open space provide the only true natural habitat refuges set
deep within urban and suburban landscapes. Such open-space habitats
function as ecological islands in a sea of urbanism. Most are necessarily
managed habitats rather than entirely natural and, like the urban environ-
ment that surrounds them, are usually subject to constant disturbance
from adjacent traffic, noise, and other forms of pollution. Economically,
since most open-space parks are set aside and maintained for a variety of
recreational purposes rather than as natural habitats, the wildlife that colo-
nizes these unnatural natural habitats must have an unusually high toler-
ance for human presence and recreational activities of all kinds.
Urban Birds
For some forms of urban wildlife, the urban landscape is merely a human-
made version of their natural environment. Thus, for pigeons the ledges,
cracks, and crevices of buildings and bridges represent an urban version of
the cracks and crevices of cliffs and rock outcrops that they use for roosting
661
and nesting in their native habitats. Similarly, the short-eared owls (Asio
flammeus) and snowy owls (Nyctea scandiaca) that show up in winter to
stand as silent sentinels at airports, golf courses, and other open areas are
simply substituting these managed short-grass habitats for the tundra hab-
itats preferred by snowy owls and the coastal marshes hunted by short-
eared owls. Their summer replacements include a host of grassland nest-
ing species such as grasshopper sparrows, kildeer, and upland sandpipers,
which all find these managed habitats to be ideal substitutes for the native
grasslands which they displaced or replaced.
Many bird inhabitants of urban and suburban environments are exotics
which were deliberately or inadvertently introduced into urban areas. Cer-
tainly the three birds with the widest urban distribution in North America,
the pigeon or rock dove, European starling, and house sparrow or English
sparrow, all fit within this category. The introduction of the European star-
ling into North American cities and suburbs resulted from the dedicated
efforts of the American Acclimitization Society of the late 1800’s. The goal
of this society was the successful introduction of all birds mentioned in the
works of Shakespeare into North America. Unfortunately for North Amer-
icans, the character of Hotspur in Henry IV makes brief note of the starling,
so the society repeatedly attempted to introduce the starling into Central
Park until they were finally successful. Since then, the starling has become
the scourge of cities and suburbs throughout much of North America and
the rest of the civilized world. Starlings damage and despoil crops, and
dirty buildings with their droppings.
The association between house sparrows and urban centers is appar-
ently very old. Evidence suggests that they abandoned their migratory
ways to become permanent occupants of some of the earliest settlements
along the Nile and Fertile Crescent, a trend that has continued to this day.
Sparrows and starlings both share certain characteristics that enable them
effectively to exploit urban and suburban habitats; both are aggressive col-
onizers and competitors, able to feed opportunistically on grains, crops,
discarded bits of garbage, and other food supplies.
Avian occupants also include an increasing diversity of released caged
pets, avian and otherwise. Thus, urban locales in Florida, Southern Califor-
nia, and along the Gulf Coast support an ever increasing diversity of para-
keets, parrots, finches, and lovebirds, all stemming from caged pet birds ei-
ther deliberately released or lost as escapees.
Feral Animals
Feral animals, mostly dogs (Canidae) and cats (Felidae), represent another
important source and component of urban wildlife. Feral dogs revert to
662
Along with squirrels, opossums, rats, coyotes, deer, bears, and other animals, the
omnivorous raccoons are among the most familiar forms of wildlife that coexist
with humans in urban and suburban neighborhoods, eating everything they can
find and often so accustomed to the human presence that they approach people to
beg for handouts. (PhotoDisc)
primal adaptive behaviors, gathering in loose packs that usually forage

and take shelter together, but have limited success because almost all cities
in developed countries have ongoing measures to control and remove
them whenever found. Feral cats are often more successful because they
are secretive, mostly nocturnal, and can better exploit available urban food
sources. The role of other feral animals as urban wildlife, mostly escaped
pets, is not well known.
Humans and Urban Wildlife

The attitude of urban dwellers toward urban wildlife varies greatly. For
many humans, urban wildlife offers a welcome respite from their other-
wise dreary and mundane surroundings. Urban wildlife in all of its forms
and colors can be aesthetically attractive, even beautiful, and is also com-
pellingly interesting. For example, the nesting of a pair of red-tailed hawks
(Buteo jamaicensis) in New York City’s Central Park sparked a remarkable
interest in bird-watching in the city and a heightened awareness of exactly
how exciting wildlife watching can be. All facets of the pair’s courtship
and nesting were observed and reported in newsprint, novellas, and even
663
a book, Red-Tails in Love. Other animals, while not nearly as large, conspic-
uous, and glamorous in their color and disposition, also elicit interest. Ur-
ban wildlife adds lively color and contrast to the otherwise monotonous
gray and grime of streets and sidewalks. Part of the attraction is that urban
birds are usually already sufficiently tolerant to be semitame in spirit, eas-
ily seen and observed, and, in some instances, easily attracted by strategi-
cally placed bird feeders and birdhouses.
Public attitudes toward urban predators vary considerably. Some peo-
ple find them attractive and interesting and even put out food for them.
Others consider them pests or potentially dangerous and avoid them. Dur-
ing rabies outbreaks or public scares, most urban wildlife is targeted by
various control programs to remove unwanted animals.
Suburban Wildlife Habitats

The vast sprawl of suburbs across the landscape offers many types of wild-
life yet another habitat to exploit, either as residences or as waystations
during the search for food or shelter. Like urban areas, suburbs offer a
range of differing habitats. The simplest suburbs are merely extensions of
urban row houses with minimal yardscapes, but there is an increasing pro-
gression toward more open and natural yards in outlying suburbs that
merge with rural areas and natural habitats. The larger and more diverse
yards at the edges of suburbs often help blur the distinction and diversity
between human landscapes and natural landscapes.
Ornamental trees, shrubs, flowers, gardens, and lawns that characterize
almost all suburban habitats provide a series of artificial habitats that can
actually increase wildlife diversity. Again, the chief wildlife benefactors
are species that can best ecologically exploit the unnatural blend of wood-
land, edge, and meadow that suburban landscapes offer. It is no accident
that some of the most common components of suburban wildlife include
thrushes such as robins, finches, cardinals, titmice, blue jays, crows, and
many other similar birds. All of these species are actually responding to the
structural components of the suburban landscape, which provide suitable
substitutes for their natural landscapes.
The blend of ornamental and garden vegetation offered by most subur-
ban landscapes offers food for a diversity of what were once consid-
ered less tolerant wildlife. Deer, wild turkey, grouse, and a host of other
animals, large and small, make periodic forays into suburbs in search
of food. Crepuscular and nocturnal wildlife is much more likely effectively
to exploit food sources offered by suburban landscapes than diurnal wild-
life, which is more at risk because of its high visibility during daylight
hours.
664
Predators
Well-wooded suburban habitats that attract a variety of wildlife also at-
tract an increasing number of predators. American kestrels (Falco spar-
verius), Cooper’s hawks (Accipiter cooperi), barn owls (Tyto alba), screech
owls (Otus spp.), and little owls (Athene noctua) provide but a small sam-
pling of birds of prey that nest deep within urban and suburban environ-
ments, taking advantage of open-space habitats deep within cities and
quickly exploiting unused areas within most suburbs. Terrestrial predators
are almost equally common, but most are nocturnal or nearly so; conse-
quently, their contacts with humans are quite limited. Many urban preda-
tors, are, in fact, mistaken for neighborhood pets and left alone or are rec-
ognized and avoided: Coyotes (Canis latrans) are often mistaken for dogs,
especially when seen in twilight. The wily coyote is equally at home in the
suburbs of Los Angeles, California, and the urban parks of New Haven,
Connecticut, joining a host of small and medium-sized mammal predators
such as foxes (Vulpes spp.) and scavengers such as opossums (Didelphis
marsupalis), raccoons (Procyon lotor), and skunks (Mephitis mephitis). These
urban predators have many behavioral attributes in common. All are om-
nivorous and able to feed on a wide variety of natural foods such as fruit,
small birds and mammals, insects, and invertebrates such as beetles, grass-
hoppers, and earthworms.
Foraging and food habits of urban predators sometimes conflict with
human concerns. Urban foxes hunt and kill cats, especially kittens, if given
the opportunity, while the larger and stronger urban coyote will often not
hesitate to kill and eat cats and dogs, to pet owners’ dismay.
Wildlife Management
Urban wildlife must be much more closely managed than wildlife of natu-
ral environments because urban and suburban habitats attract an enor-
mous number of pest species as well as interesting and beneficial species.
Introduced species such as starlings may also transmit histoplasmosis, a
fungal disease that attacks human lungs. Other birds may also be harbin-
gers, carriers, and vectors of various diseases, the most notable of which
are the parrots and parakeets, which transmit parrot fever or psittocosis.
Rats and mice (Rodentia) carry and spread disease and despoil both resi-
dential and public buildings and other structures.
The growing interest in urban wildlife has stimulated innumerable pro-
grams to promote beneficial wildlife. Both public and private organiza-
tions and agencies have embarked on a variety of programs aimed at re-
modeling existing habitats and even creating new habitats for urban
wildlife.
665
Programs aimed at creating new or modifying existing urban habitats

come in a variety of categories, such as linear parks, greenways, urban
wildlife acres programs, backyard gardens, and treescaping streets and
roadways, all of which create biodiversity, which in turn provides attrac-
tive habitats for colonization by additional animals and plants. Modifica-
tions of existing habitats to increase animal biodiversity include “critter
crossings,” roadside habitats, backyard gardens, arbor plantings, all of
which provide refuges, shelters, breeding sites, connecting corridors, and
safe havens that promote the welfare of urban and suburban wildlife.
Many existing open-space habitats are also being modified. Many ur-
ban renewal commissions have placed new and more restrictive regu-
lations on the use of pesticides and fertilizers on golf courses, which not
only reduces incidence and intensity of nonpoint pollution from the golf
courses but also reduces the incidence of wildlife poisoning. These steps
cannot help but increase the biotic potential of golf courses for supporting
local biodiversity.
Dwight G. Smith
See also: Biodiversity; Food chains and webs; Landscape ecology; Trophic
levels and ecological niches; Wildlife management.

Adams, Lowell W. Urban Wildlife Habitats: A Landscape Perspective. Minne-
apolis: University of Minnesota Press, 1994.
Bird, David, Daniel Varland, and Juan Josè Negro, eds. Raptors in Human
Landscapes. San Diego, Calif.: Academic Press, 1996.
Forman, Richard, and Michel Godron. Landscape Ecology. New York: John
Wiley & Sons, 1986.
Gill, Don, and Penelope Bonnett. Nature in the Urban Landscape: A Study of
City Ecosystems. Baltimore: York Press, 1973.
McDonnell, Mark J., and Steward T. A. Pickett, eds. Humans as Components
of Ecosystems. New York: Springer-Verlag, 1993.
666
WASTE MANAGEMENT
Types of ecology: Ecotoxicology; Restoration and conservation ecology
Waste management concerns the physical by-products of human activity that can-
not be reintegrated into the ecological biomass cycle. These by-products include
solid, liquid, and airborne substances that are potentially harmful to living organ-
isms. As the human population grows and the use of manufactured materials ex-
pands, disposing of waste becomes more challenging.
A ccording to the World Watch Institute, world production of manufac-

tured materials (not counting recycled materials) increased nearly
2.5 times between the early 1960’s and the late 1990’s. In industrialized
countries the increase is far greater: The United States, for example, has
seen an eighteenfold increase in materials production since 1900. The aver-
age U.S. citizen throws away an estimated 2 to 8 pounds of garbage daily,
and although studies demonstrate that the per-person production of waste
remained approximately the same throughout the twentieth century, the
sharp rise in population and expanding industrial base meant greater total
accumulations of waste. Furthermore, the types of waste changed.
Waste is commonly categorized as domestic, or solid, waste, and indus-
trial, or liquid, waste, although the distinction is not absolute. Both may
contain toxic substances, but the percentage of toxins in industrial waste is
likely to be higher, and the types of waste are disposed in different ways.
The smoke emitted from industrial processing of materials and vehicle ex-
haust are additional types of waste, although they are commonly thought
of as pollution rather than waste.
Solid Waste
Solid waste is the familiar garbage that households and businesses in the
United States have sent to the dump since garbage collection began late in
the nineteenth century. The largest portion, more than 40 percent, consists
of paper products, especially newspaper and containers. Yard waste, food
debris, plastic containers and wrappings, bottles, metals, and appliances
are also regularly thrown away. About 1 percent of this waste involves haz-
ardous materials, typically insecticides, beauty aids, and cleaning prod-
ucts. Construction waste accounts for a large share—about 12 percent—of
solid waste and may contribute a higher proportion of hazardous materi-
als, such as solvents and paint.
Although most of these materials are solid, when dumped together
667
Waste management
A wastewater treatment plant

collects the domestic and
industrial effluent that has
been conveyed to its location
by a sewage system and treats
this contaminated water by
removing solids, filters,
biological decomposition, and
other processes, ending in
release into the ground or,
more usually, into a surface
watercourse. (PhotoDisc)
they can soak up rainwater and then ooze chemical-laden liquids. This
leachate may filter down into the groundwater and pollute nearby streams
and wells. If it contains toxic elements, such as the lead or mercury from
batteries, the leachate can be dangerous to health. The odor from rotting
garbage may also foul the air, seldom enough to be harmful but still repel-
lent to people living nearby. It can attract animal scavengers, which may
become infected with diseases from the garbage and spread them to other
animals or even humans, especially if feces are part of the waste.
In order to combat these effects, sanitary landfills place a plastic lining
under the waste to contain leachate and cover each day’s load of garbage
under a thin layer of soil. Pipe systems also disperse methane gas pro-
duced by rotting organic materials. The landfills are therefore less danger-
ous to human health or the environment, but many old, abandoned sites
were not so well engineered. They may continue to dribble harmful chemi-
cals into groundwater for decades and emit methane, which is flammable.
Numerous small, illegal dumps and litter compound the problem.
Measures to reduce the amount of waste deposited in landfills have
partially succeeded. Recycling has drastically cut the total paper, metal,
668
Waste management
and glass waste in some U.S. states and industrialized countries. The use of
garbage disposals and composting has caused the proportion of organic
materials to decline. However, such reductions did not eliminate solid
waste. By the end of the twentieth century, cities were finding it increas-
ingly difficult to find room for new landfill sites, even when the space was
urgently needed. Stringent regulations about the geological composition
of landfills reduced the number of usable sites, while objections from citi-
zen action committees, known as “not in my back yard” (NIMBY) groups,
also eliminated sites near populated areas.
Facilities used to incinerate waste, which sometimes powered electrical
generators with the resulting heat energy, also faced objections because
burning could release health-threatening materials, such as dioxins, into the
air. Moreover, a significant proportion of waste, such as appliances and con-
crete, cannot be eliminated by burning. Tires, too hazardous to burn, float
to the surface in landfills, causing continuous problems for waste manag-
ers; they often end up stacking the tires in immense piles that, if acciden-
tally ignited, can burn out of control and create large clouds of black fumes.
Industrial Waste
The effluent stream of by-products from factories, as well as chemical and
petroleum refineries, is made up of water, solid filings and cuttings, liquid
solvents and oil derivatives, and semisolid sludge. The solid components
are usually no more hazardous than household wastes, although medical
waste—particularly tainted blood and used “sharps,” such as needles and
scalpels—may pose the additional danger of spreading disease. However,
liquids and semiliquids sometimes contain a high proportion of hazardous
chemicals. Rain also leaches chemicals, such as cyanide and mercury, out
of the smelted tailings from mines. Agricultural fertilizers and pesticides
can enter groundwater or streams as well. Because these liquid wastes rap-
idly spread through waterways and groundwater, they are often collec-
tively known as toxic waste.
Industry now uses all ninety-two naturally occurring elements on the
periodic table, and the isotopes of some of these are radioactive. Nuclear
weapons manufacturing in particular leaves radioactive debris, but medi-
cal procedures that use radioactive tracers and scientific instruments may
also create radioactive wastes. This nuclear waste continues to emit radia-
tion for thousands or hundreds of thousands of years, and improperly
stored radioactive materials have been associated with increased risk of
disease for people, animals, and plants.
During the 1980’s and 1990’s federal and state regulations brought in-
dustrial waste management under rigorous control. Facilities known as se-
669
Waste management
cure landfills are designed to contain nonradioactive industrial wastes in

tightly lined, self-contained areas. Incinerators reduce the waste to harm-
less ash while releasing few or no harmful particulates into the atmo-
sphere. Separate repositories store nuclear wastes deep underground in
leak-proof containers.
The public is seldom reassured by such measures, however. Leakage oc-
casionally occurs from secure landfills. Near-zero toxic emissions from in-
cineration means that some toxins do, in fact, escape into the atmosphere.
In addition, nuclear repositories may not be catastrophe proof; for exam-
ple, an earthquake could crack open containers, releasing radioactive ma-
terial into groundwater supplies. Although waste managers insist that
these dangers are minimal, the news media bring them to public attention,
and NIMBYs regularly resist the opening of new secure landfills and radio-
active waste repositories. State governments often object as well, as was
the case when the Nevada legislature stalled the construction of a nuclear
repository at Yucca Mountain. Many old facilities, built before strict gov-
ernment oversight, remain in use and could leak toxic materials into the
environment undetected. The memory of deadly chemical leaks, such as
that discovered at Love Canal in New York in 1976, and of released radio-
active material, such as the plutonium that escaped the Hanford Nuclear
Reservation in Washington State, makes the public wary of hazardous
wastes.
As a result of citizen concern, most new hazardous waste disposal sites
are now located far from population centers. This has created a new peril.
The waste must be transported, primarily by trucks and trains, to a facility.
Traffic accidents and train derailings en route can dump extremely danger-
ous chemicals straight into water or the atmosphere. Evacuations of resi-
dents near such accidents, while not common, increased during the 1990’s.
Even if people are rescued, however, plant and animal life is not safe-
guarded.
Environmental Consequences
Many critics of waste management insist that only source reduction—a
drastic decrease in the use of raw materials—will make waste disposal
safe. Accordingly, during the 1990’s some countries, notably Denmark and
Germany, sought to reduce virgin material use as much as 90 percent by in-
tensifying recycling. In the United States, Superfund legislation was
passed to set aside federal funds to pay for cleanups of the most dangerous
hazardous waste sites. Other industrial countries have similar projects.
Still, only a fraction of sites receive attention, and until source reduction
goals are met, household and industrial wastes will continue to swell land-
670
Waste management
fills with environmentally hazardous substances. Illegal dumping of haz-

ardous waste exacerbates the danger.
Scientists disagree about how severely wastes damage the environ-
ment, but there is agreement that repercussions are evident and likely to
increase. Methane from dumps, smoke-stack emissions, and vehicle ex-
haust contain greenhouse gases, which are implicated in global warming.
Nutrients released from sewers, as well as runoff from agriculture and
mining, degrade the environment of rivers and streams, harming aquatic
life and leaving the water unusable without special treatment. The
waterborne wastes that reach the ocean, supplemented by ocean dumping
of toxic materials, alter and sometimes destroy offshore ecosystems, as is
the case for many coral reefs worldwide.
Roger Smith
See also: Biological invasions; Biomagnification; Biopesticides; Deforesta-

tion; Eutrophication; Genetically modified foods; Integrated pest manage-
ment; Invasive plants; Ocean pollution and oil spills; Ozone depletion and
ozone holes; Pesticides; Phytoplankton; Pollution effects; Slash-and-burn
agriculture.

Baarschers, William H. Eco-Facts and Eco-Fiction: Understanding the Environ-
mental Debate. New York: Routledge, 1996.
Dunne, Thomas, and Luna B. Leopold. Water in Environmental Planning.
San Francisco: W. H. Freeman, 1978.
Gourlay, K. A. World of Waste: Dilemmas of Industrial Development. New
York: St. Martin’s Press, 1992.
Laak, Rein. Wastewater Engineering Design for Unsewered Areas. Lancaster,
Pa.: Technomic, 1986.
McGhee, Terrence. Water Supply and Sewerage. New York: McGraw-Hill,
1991.
Qasim, Syed R. Wastewater Treatment Plants: Planning, Design, and Opera-
tion. 2d ed. Lancaster, Pa.: Technomic, 1999.
Rathje, William, and Cullen Murphy. Rubbish! The Archaeology of Garbage.
Tucson: University of Arizona Press, 2001.
Salvato, Joseph A. Environmental Engineering and Sanitation. 4th ed. New
York: Wiley, 1992.
Tillman, Glenn M. Wastewater Operations: Troubleshooting and Problem
Solving. Chelsea, Mich.: Ann Arbor Press, 1996.
Whitaker, Jennifer Seymour. Salvaging the Land of Plenty: Garbage and the
American Dream. New York: William Morrow, 1994.
671
WETLANDS
Wetlands, transitional areas between aquatic and terrestrial habitats, are home to a
variety of flood-tolerant and salt-tolerant plant species.
W etlands represent one of the most biologically unique and produc-

tive of all natural habitats. In their unaltered state, these water-
influenced areas are used by a variety of wildlife species. These habitats
also have the ability to take up and store water during floods, and their
soils and plants have the ability to remove nutrients and heavy metals
from water. The recognition of these values helped to slow the rate of
wetlands loss to such uses as agricultural development and urban expan-
sion. A desire to protect remaining wetland acres has led to a significant
movement for wetlands preservation.
Definition of Wetlands
Ecologists recognize wetlands as a type of ecotone. Ecotones are unique ar-
eas that represent a transition from one type of habitat to another. Often,
these transitional areas have characteristics of both habitats. Wetlands
are areas located between aquatic, water-based habitats and dry land. Be-
cause they are located at the edge of an aquatic habitat, wetlands are al-
ways influenced to some degree by water. They are not always under
water, as are aquatic habitats, and they are not always dry, as are terrestrial
habitats.
The most important environmental factor in wetlands is the periodic or
frequent occurrence of water. This presence of water influences both the
nature of the soil and the flora and fauna of a region. Soils which experi-
ence periodic coverage with water become anoxic, develop a dark color,
and give off an odor of hydrogen sulfide. These soil characteristics differ
from those of upland soils and give wetland soils their unique hydric na-
ture. In these soils influenced by water, only flood-tolerant hydrophyte
species can exist. Hydrophytic plants vary in their tolerance to flooding
from frequent (such as bald cypress) to infrequent (such as willows).
In defining a particular area as a wetland, often all three of the compo-
nents listed above are used: water, hydric soils, and hydrophytic plants.
However, the presence of water is not always a reliable indicator because
water rarely covers a wetland at all times. Often, a wetland is dry during a
period of low river flow or during a low tide. For this reason, only hydric
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Wetlands
soils and hydrophytic plants should be used as reliable indicators of a wet-

land.
The broadest classification of wetlands includes two categories: fresh-
water and saltwater wetlands. Freshwater wetlands occur inland at the
edges of rivers, streams, lakes, and other depressions that regularly fill
with rainwater. Saltwater wetlands occur along the coast in bays, where
salt water and fresh water mix and wave energy is reduced.
Freshwater Wetlands
Of the two categories, freshwater wetlands are by far the most common.
Freshwater wetlands are subdivided into two categories: tree-dominated
types and grass-dominated types. Tree-dominated freshwater wetlands
include areas that are frequently covered with water (such as cypress
swamps) and those that are only occasionally covered with water (such as
bottomland forests). Grass-dominated types include freshwater marshes,
prairie potholes, and bogs.
While freshwater marshes are widespread, prairie potholes and bogs
occur regionally in the United States. Prairie potholes are located in the
central portion of the United States, while bogs are found in the Northeast
and Great Lakes regions.
Wetlands are a special ecotone, or transitional environment, with characteristics of

both dry and aquatic habitats and therefore are home to a large variety of plants,
animals, birds, and other forms of life. (PhotoDisc)
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Wetlands
Saltwater Wetlands
Saltwater wetlands are also subdivided into tree-dominated and grass-
dominated types. Tree-dominated types include tropical mangrove swamps.
Grass-dominated types can be further subdivided into salt marshes and
brackish marshes. Salt marshes occur in bays along the coast where salt
water and fresh water mix in almost equal proportions. Brackish marshes
occur farther inland than salt marshes do; their mix contains less seawater
and more fresh water. Both grass-dominated types are common in bays
along the Gulf of Mexico and the East Coast of the United States.
The Biota of Wetlands

The most noticeable feature of all wetlands is the abundance of plant life. A
variety of plant species thrive in wetlands, but each occurs only in a partic-
ular kind of habitat. Freshwater wetlands that are frequently flooded pro-
vide a favorable habitat for water-tolerant trees, such as bald cypress and
water tupelo, and water-tolerant herbaceous plants, such as cattail, arrow-
head, bulrush, spike rush, water lily, and duckweed. Less frequently
flooded freshwater areas support trees such as willow, cottonwood, water
oak, water hickory, and red maple. Seawater areas in tropical bays favor
the development of mangroves, while temperate bays favor the develop-
ment of cordgrass.
Wetland plants provide a habitat for a variety of animals. Cypress
swamps and cattail marshes support a large assortment of animals, includ-
ing alligators, ducks, crayfish, turtles, fish, frogs, muskrat, wading birds,
and snakes. Likewise, mangrove prop roots provide attachment sites for a
variety of invertebrates and shelter for numerous small fish, while upper
branches provide roosting and nesting sites for birds. In salt marshes, mus-
sels live among cordgrass roots, while snails, fiddler crabs, oysters, and
clapper rails live among plant stalks. When water covers cordgrass at high
tide, plant stalks shelter small fish, crabs, and shrimp seeking refuge from
large predators.
The Value of Wetlands

The amount of plant material produced in wetlands is higher than that
produced in most aquatic and terrestrial habitats. This large amount of
plant material supports an abundance of animal life, including commer-
cially important species such as crayfish, ducks, fish, muskrat, shrimp, and
crabs.
The biotic value of wetlands is well recognized, but it represents only a
part of their total value. Wetlands provide “services” for other areas that
often go unrecognized. For example, freshwater wetlands are capable of
674
Wetlands
storing large amounts of water during periods of heavy rainfall. This capa-
bility can be important in minimizing the impact of flooding downstream.
Saltwater wetlands along coastlines are an effective barrier against storms
and hurricanes. These natural barriers hold back the force of winds, waves,
and storm surges while protecting inland areas. Wetlands are also capable
of increasing water quality through the trapping of sediment, uptake of
nutrients, and retention of heavy metals. Sediment trapping occurs when
moving water is slowed enough by grass and trees to allow suspended
sediment particles to settle. Wetland plants take up nutrients, such as ni-
trates and phosphates, from agricultural runoff and sewage. For this rea-
son, wetlands are used as a final treatment step for domestic sewage from
some small cities. Wetland soils are capable of binding heavy metals, effec-
tively removing these toxic materials from the water.
Wetlands Loss and Preservation

It is estimated that the United States once contained more than 200 million
acres of wetlands. Less than half this amount remains today. Once consid-
ered wastelands, wetlands were prime targets for “improvement.” Exten-
sive areas of freshwater wetlands and prairie potholes have been drained
and filled for agricultural development. Saltwater wetlands have been re-
placed by urban or residential development and covered with dredge
spoil. Wetlands loss rates have slowed, but an estimated 300,000 acres con-
tinue to be lost each year in the United States. The loss of wetlands habitat
threatens the survival of a number of animal species, including the whoop-
ing crane, American crocodile, Florida panther, manatee, Houston toad,
snail kite, and wood stork.
Since the 1970’s the rate of wetlands loss has slowed for several reasons.
One is the passage of federal and state laws designed to protect wetlands;
another is the efforts of conservation organizations. At the federal level,
the single most effective tool for wetlands preservation is Section 404 of the
Clean Water Act. Section 404 requires that a permit be issued before the re-
lease of dredge or fill material into U.S. waters, including wetlands. At the
state level, Section 401 of the Clean Water Act allows states to restrict the
release of dredge or fill material into wetlands. Subsequent legislation, no-
tably the North American Wetlands Conservation Act of 1989, worked to
conserve wetland habitat. The 1989 act was passed in part to support the
North American Waterfowl Management Plan, an international agreement
between Canada, Mexico, and the United States to protect wetland/up-
land habitats on which waterfowl and other migratory birds in North
America depend. In December, 2002, President George W. Bush signed the
North American Wetlands Conservation Reauthorization Act, intended to
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Wetlands
“keep our water clean and help provide habitat for hundreds of species of
wildlife.” Several conservation organizations also support wetlands pres-
ervation, including Ducks Unlimited, the National Audubon Society, the
National Wildlife Federation, and the Nature Conservancy. These organi-
zations keep the public informed regarding wetlands issues and are active
in wetlands acquisition.
Steve K. Alexander, updated by Christina J. Moose

forests; Rain forests; Rain forests and the atmosphere; Rangeland; Reefs;
Savannas and deciduous tropical forests; Taiga; Tundra and high-altitude
biomes.

Hey, Donald L., and Nancy S. Philippi. A Case for Wetland Restoration. New
York: Wiley, 1999.
Littlehales, Bates, and William Niering. Wetlands of North America. Char-
lottesville, Va.: Thomasson-Grant, 1991.
Mitchell, John. “Our Disappearing Wetlands.” National Geographic 182,
no. 4 (1992).
Mitsch, William J., and James G. Gosselink. Wetlands. 3d ed. New York:
John Wiley, 2000.
Monks, Vicki. “The Beauty of Wetlands.” National Wildlife 34, no. 4 (1996).
Vileisis, Ann. Discovering the Unknown Landscape: A History of America’s
Wetlands. Washington, D.C.: Island Press, 1999.
Watzin, Mary, and James Gosselink. The Fragile Fringe: Coastal Wetlands of
the Continental Unites States. Rockville, Md.: National Oceanic and At-
mospheric Administration, 1992.
676
WILDLIFE MANAGEMENT
Wildlife management strives to allow the use of ecological communities for human
benefit while preserving their ecological components unharmed. It also seeks to re-
store biological communities by managing habitats and controlling the taking of
organisms for sport or economic gain.
W ildlife management, also known as game management, is often

compared with farming or forestry, because one of its goals is to
ensure annual “crops” of wild animals. Conservationist Aldo Leopold, in
1933, defined “game management” as the art of making land produce
sustained annual crops of wild game for recreational use. At that time,
animals considered to be game included deer and animals such as coy-
otes that do damage to domestic animals or crops. Now, however, the
term “wildlife” has replaced game, and virtually all living organisms, in-
cluding invertebrates and plants, are included in management consider-
ations.
Approaches to Wildlife Management

The process of wildlife management has moved through a sequence of six
approaches: the restriction of harvest (by law); predator control; the estab-
lishment of refuges, reserves, and parks; the artificial stocking of native
species and introduction of exotic ones; environmental controls, or man-
agement of habitat; and education of the general public. All six are used in
modern wildlife management programs, but most emphasis is placed on
habitat management and control of harvest.
All fifty states of the United States have departments responsible for
wildlife conservation. An appointed board of directors or commission
oversees the actions of the departments. Groups for wildlife law enforce-
ment, research, management, and information and education make recom-
mendations to the board of directors regarding wildlife management ac-
tions. The federal government of the United States also has many agencies
that manage wildlife on public lands. The U.S. Fish and Wildlife Service is
involved with animals that cross state lines, including migratory birds
such as waterfowl, marine mammals, and any plants and animals listed as
rare or endangered under the National Environmental Protection Act of
1970. Other agencies, such as the U.S. Forest Service, Bureau of Land Man-
agement, Soil Conservation Service, and the U.S. National Park Service, do
677
Wildlife management
extensive wildlife work. Many private organizations, such as the National

Wildlife Federation, the Audubon Society, and the Sierra Club actively pro-
mote wildlife conservation.
Wildlife management decisions involve the entire range of biological,
sociological, political, and economic considerations of human society. To-
day, the wildlife resource in the United States is managed primarily either
for consumptive use (such as sport hunting) or for nonconsumptive use
(such as bird-watching). Virtually all wildlife management problems are
related to the large human population of the earth. Some specific problems
are habitat loss (for example, the destruction of tropical rain forests), pollu-
tion, diseases introduced by domestic animals into wildlife populations,
and the illegal killing of animals for their parts, such as the poaching of ele-
phants for their ivory.
Managing Wildlife Communities

A wildlife manager must first determine the physical and biological condi-
tions of the organism or organisms being managed. Issues include what
the best habitat for the animal is and how many animals this habitat can
support. The stage of ecological succession determines the presence or ab-
sence of particular animals in an area. All animals need food, water, and
protection from weather and predators. Special needs, such as a hollow
tree in which to raise young, for example, must be fulfilled within the ani-
mal’s home range. Wildlife managers attempt to remove or provide items
that are most limiting to a population of animals. In many respects, solving
wildlife management problems is an art; it is similar to medicine in that it
often must deal with symptoms (birds dying, for example) and imprecise
information.
The stage of ecological succession may be maintained by plowing
lands, spraying unwanted plants with a chemical to kill them, or using fire,
under controlled conditions, to burn an area to improve the habitat for a
certain wildlife group. Refuges and preserves may be set aside to assure
that some of the needed habitat is available; nest boxes and water supplies
may even be provided.
Periodic surveys of the number of animals in a population provide
guidelines for their protection. If animals are more abundant than the low-
est carrying capacity, a controlled harvest may be allowed. Sustained an-
nual yield assures that no more than the population surplus is taken. Wild-
life laws protect the animals, provide for public safety, often set ethical
guides for sporting harvest, and attempt to provide all hunters with an eq-
uitable chance of obtaining certain animals (for example, by setting bag
limits). If proper wildlife management procedures are followed, no animal
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Wildlife management
need become rare or endangered by sport hunting. Market hunting, the

taking of animals for the sale of their products, such as meat or hides, has
been stopped in the United States since the 1920’s and is also illegal in most
other areas of the world. There are almost no societies left that are true sub-
sistence hunters—that is, living exclusively on the materials produced by
the wildlife resource.
The Need for Wildlife Management

The proper management of wildlife resources, based on sound ecological
principles, is essential to the well-being of humans. All domestic plants
and animals came from wild stock, and this genetic reservoir must be
maintained. Maintaining the web of life that includes these organisms is
necessary for human survival. Wildlife resources are used by at least 60
percent of the citizens of the United States each year, and about 6 percent
are sport hunters. Wildlife provides considerable commercial value from
products, such as meat; it also offers aesthetic values of immeasurable
worth. Seeking and observing wildlife provides needed relief from the ev-
eryday tensions of human life. Moreover, by observing wildlife reactions
to environmental quality, investigators can monitor the status of the bio-
logical system within which humans live. Wildlife populations serve as a
crucial index of environmental quality.
Perhaps most important of all, wildlife management helps preserve the
biodiversity of communities and ecosystems. Without an effort toward in-
cluding these ecological considerations along with economic resource
concerns, habitat and species loss would quickly ensue, even more rapidly
than it now does, in response to urban, suburban, industrial, and agri-
cultural development. Wildlife management is a dynamic activity that,
to be effective, must reflect an understanding of and respect for the natu-
ral world. It cannot be practiced in a vacuum but must encompass the
realm of complex human interactions that often have conflicting goals and
values. Aldo Leopold once defined conservation as man living in harmony
with the land; successful wildlife management will help assure that this oc-
curs.
David L. Chesemore

animal species; Endangered plant species; Erosion and erosion control;
Forest management; Genetic diversity; Grazing and overgrazing; Habitats
and biomes; Landscape ecology; Multiple-use approach; Old-growth for-
ests; Reforestation; Restoration ecology; Species loss; Sustainable develop-
ment; Urban and suburban wildlife; Zoos.
679
Wildlife management

Anderson, S. H. Managing Our Wildlife Resources. 3d ed. Upper Saddle
Bailey, James A. Principles of Wildlife Management. New York: John Wiley &
Sons, 1984.
Bissonette, John A., ed. Wildlife and Landscape Ecology: Effects of Pattern and
Scale. New York: Springer, 1997.
Cooperrider, Allen Y., R. J. Boyd, H. R. Stuart, and Shirley L. McCulloch. In-
ventory and Monitoring of Wildlife Habitat. Washington, D.C.: U.S. Gov-
ernment Printing Office, 1986.
Dasmann, R. F. Wildlife Biology. New York: John Wiley & Sons, 1981.
Di Silvestro, Roger, ed. Audubon Wildlife Report, 1986. New York: National
Audubon Society, 1986.
Giles, R. H., Jr. Wildlife Management. San Francisco: W. H. Freeman, 1978.
Leopold, Aldo. Game Management. New York: Charles Scribner’s Sons,
1939. Reprint. Madison: University of Wisconsin Press, 1986.
Matthiessen, Peter. Wildlife in America. New York: Viking Press, 1987.
Robinson, W. L., and E. G. Bolen. Wildlife Ecology and Management. 4th ed.
Upper Saddle River, N.J.: Prentice Hall, 1999.
680
ZOOS
Keeping wild animals has evolved, over the past five thousand years, from animal
collections maintained by ancient societies to modern zoological gardens and
aquariums with significant programs in wildlife appreciation, education, science,
and conservation. Originally entertainment venues, zoos have shifted their focus
to education and active conservation of endangered and threatened species.
N early 600 million people worldwide visit a zoo each year—roughly

10 percent of the global population. Modern zoos, often called wild-
life conservation parks or natural wildlife parks, have replaced cages of
concrete and steel with simulated natural environments, and new animals
are obtained via selective breeding instead of being captured from the
wild.
History of Zoos
Early zoos were the sole province of the wealthy; the first recorded zoo
in history belonged to a Chinese emperor in 1100 b.c.e. It was not until the
nineteenth century that zoos were open to the public. The word “zoo” de-
rives from the phrase “zoological park,” and that was what the first zoos
were designed to be: afternoon diversions along the same lines as the
amusement park or the circus. Exotic beasts from newly charted regions
were captured and displayed with little regard for their health or emo-
tional well being. Mortality was high, and display animals were constantly
replaced with animals captured from the wild, of which there seemed to be
an inexhaustible supply.
The first zoo to use moats to separate animals from visitors was estab-
lished in Germany by Carl Hagenbeck in 1907. These moats provided visi-
tors with an unobstructed view and, depending on their placement, made
it seem as if the animals were free. While the bars were gone, the habitat
was still nothing like what the animals were accustomed to in the wild.
Those animals that did not spend their days sleeping often displayed near-
psychotic behavior patterns, such as pacing, head butting, and even self-
mutilation.
A New Role
Two things changed the way zoos functioned during the twentieth cen-
681
Zoos
tury. First, movies and television allowed potential visitors to see the ani-
mals in their natural habitats, and suddenly giraffes, lions, and zebras
were no longer quite so exotic. Second, wild animals were becoming more
scarce, and words such as “conservation” and “endangered” entered the
collective vocabulary. Acquiring specimens from the wilderness became
more costly, and zoos began to look at internal breeding programs to re-
plenish their stock. However, they found that animals kept in unnatural
and in some cases inhumane conditions would not breed.
New zoo enclosures were designed to encourage natural behavior in
animals by replicating their natural environment as much as possible
while still ensuring the safety of both the animals and the zoo visitors. Ani-
mals began receiving healthier diets and, when possible, were allowed to
feed in much the same way they would in the wild—by digging, foraging,
or grazing. Human contact with orphaned and injured animals was kept to
an absolute minimum, and some zoos took the additional step of not nam-
ing their animals to discourage anthropomorphism. By 1995, 80 percent of
the mammals on display in zoos were born in captivity.
Image Not Available
682
Zoos
Another trend that began in the late twentieth century was the building
of “wild animal parks.” The San Diego Zoo, for example, established one
of the earliest wild animal parks northeast of San Diego near Escondido,
California, which turns the tables by restricting human visitors behind
fences and within tram cars as the animals roam freely over large land
tracts that approximate their natural habitats.
Mission: Preservation
As concern over endangered animal species, coupled with ecologists’
alarm over a decline in biodiversity, spread at the end of the twentieth
century, zoos increasingly took on the mission of preservation and breed-
ing. The American Zoo and Aquarium Association (AZA), a nonprofit
organization dedicated to the advancement of zoos and aquariums in
the areas of conservation, education, science, and recreation, states its
members’ mission as “work[ing] cooperatively to save and protect the
wonders of the living natural world.” The AZA has accredited more
than two hundred zoos worldwide and sponsors a variety of research, ed-
ucation, and preservation programs. One of these, the Species Survival
Plan (SSP) program, was established in 1981 to “help ensure the survival
of selected wildlife species into the future and to provide a link be-
tween zoo and aquarium animals and the conservation of their wild coun-
terparts” through breeding and, where possible, reintroduction into the
wild.
Zoo managers continue to struggle to balance science, conservation bi-
ology, scarce resource allocation, and ethics. Among the choices that must
be made are whether predators should be offered the chance to exercise
natural hunting behaviors by being offered live prey, or whether zoos
should maintain potentially deadly animals that are necessary for breed-
ing programs but are dangerous and difficult to control, such as macaques,
many of which harbor the deadly hepatitis B virus, or adult male ele-
phants. Another dilemma is the question of what should become of “sur-
plus” animals that are inbred, unable to reproduce, or are otherwise geneti-
cally inferior.
Municipal bureaucracies can also hamper zoo conservation efforts. Zoo
managers must often combat local governments and public opinion when
dealing with unpopular issues, such as surplus animals and resource allo-
cation. In addition, budget cuts have forced zoo managers to turn to the
private sector for financial assistance. Fund-raising activities range from
the traditional “adopt an animal” programs to the extraordinary commer-
cial venture of selling “exotic compost.”
There are those who question whether zoos should exist at all—
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Zoos
whether it is cruel or unusual to take animals from their natural habi-

tat and place them on display. The People’s Republic of China, for ex-
ample, has “rented” its zoo’s giant pandas when the pandas might be
better served by remaining in the wild or in a captive-breeding pro-
gram that would allow them to replenish their numbers. Critics claim that
the money devoted to zoos and captive-breeding programs would be
better spent on preserving the animals’ natural habitats. To combat these
types of criticism, some zoos began to change their focus from “collect-
ing” wildlife to “protecting” wildlife, also known as field conservation.
In these new exhibits, zoo visitors view exhibits linked with protection and
conservation programs in natural habitats, allowing visitors to connect
what they are seeing in captivity to what’s worth saving in the wilder-
ness. Some zoos have taken the additional step of “adopting” wildlife ref-
uges.
Despite the criticism, the fact remains that there are many animal spe-
cies that could not survive without the existence of zoos and captive-
breeding programs. Ironically, where historical zoos replenished their
stock from the wilderness, some zoos are now replenishing the wilderness
with captive-bred animals.
P. S. Ramsey, updated by Christina J. Moose

animal species; Genetic diversity; Restoration ecology; Species loss; Sus-
tainable development; Urban and suburban wildlife; Wildlife manage-
ment.

Bell, Catharine, et al., eds. Encyclopedia of the World’s Zoos. Chicago: Fitzroy
Dearborn, 2001.
Croke, Vicki. The Modern Ark: The Story of Zoos, Past, Present, and Future.
New York: Charles Scribner’s Sons, 1997.
Hoage, R. J., and William A. Deiss, eds. New Worlds, New Animals: From Me-
nagerie to Zoological Park in the Nineteenth Century. Baltimore: The Johns
Hopkins University Press, 1996.
Kisling, Vernon N., Jr., ed. Zoo and Aquarium History: Ancient Animal Collec-
tions to Zoological Gardens. Boca Raton, Fla.: CRC Press, 2001.
Koebner, Linda. Zoo Book: The Evolution of Wildlife Conservation Centers.
Preface by William Conway. New York: T. Doherty, 1994.
Norton, Bryan G., Michael Hutchins, Elizabeth F. Stevens, and Terry L. Ma-
ple, eds. Ethics on the Ark: Zoos, Animal Welfare, and Wildlife Conservation.
Washington, D.C.: Smithsonian Institution Press, 1995.
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Zoos
Tudge, Colin. Last Animals at the Zoo: How Mass Extinction Can Be Stopped.
Washington, D.C.: Island Press, 1992.
Wemmer, Christen M., ed. The Ark Evolving: Zoos and Aquariums in Transi-
tion. Front Royal, Va.: Smithsonian Institution Conservation and Re-
search Center, 1995.
685
GLOSSARY
abiotic: Not living; used to refer to the nonliving elements of an ecosystem
or biome, consisting of climate, the minerals in soil, rocks, water, oxy-
gen, carbon dioxide, and other physical components.
abundance: The density and prevalence of organisms living in a particular
population, community, or ecosystem.
abyssal marine zone: The dark marine zone that extends from the ocean
floor to where the continental slope begins.
acclimatization: A process by which animals habituate their physiological
responses to the conditions of a particular environment.
acid deposition: The process whereby acidic (very low pH, usually under
5.6) gases, particles, and precipitation (rain, fog, dew, snow, or sleet) fall
on the surface of the earth.
active or wide foraging: Moving across a relatively large area in search of
prey.
adaptation: In evolutionary biology, a heritable structure, physiological
process, or behavioral pattern that gives an organism a better chance of
surviving and reproducing; in physiology, the change in the response
of a sense organ following continuous application of a constant stim-
ulus.
adaptive radiation: The relatively rapid evolution of several new species
from a common ancestor following invasion of a new geographic region
or ecological niche, or exploitation of a new ecological opportunity.
aerobic: Characterizing any biological process that occurs in the presence
of free oxygen.
aestivation: See estivation.
aggregation: A group of organisms that live in closer proximity than they
would were they to be randomly or evenly distributed. Aggregations of
the same species are known as populations.
aggression: A physical act or threat of action by one individual that re-
duces the freedom or genetic fitness of another.
agricultural ecology: Also called agroecology, the study of agricultural eco-
systems, their components (such as crop species), functions, interactions,
and impact on natural ecosystems and abiotic factors such as atmo-
spheric and water systems—often with an emphasis on the develop-
ment of sustainable systems.
agricultural revolution: The transition by humans from hunting and gath-
ering all their food to domesticating plants for food.
alleles: Different forms of a particular gene, located at the same position on
687
Glossary
a chromosome. Most genes have at least two naturally occurring alleles.

These alleles may be the same or may be different.
allelochemical: A general term for a chemical used as a messenger be-
tween members of different species; allomones and kairomones are al-
lelochemicals, but hormones and pheromones are not.
allelopathy: The phenomenon in which an organism produces and releases
chemicals that are toxic to, or inhibit the growth of, another organism,
although sometimes with beneficial effects to the other organism.
allopatric speciation: The evolution of two new species as a result of the
separation of two groups of the same species from each other.
allopolyploidy: Formation of a new species resulting from the mating of
two different species, resulting in a sterile or reproductively isolated hy-
brid.
alpine tundra: The area of Europe, Asia, and North America north of the
boreal coniferous forest zone, where the soils remain frozen most of the
year, constituting about 3 percent of the earth’s surface.
altruism: A behavior that increases the fitness of the recipient individual
while decreasing the fitness of the performing individual.
amensalism: An interspecies relationship in which one species is harmed
while the other remains uneffected.
anaerobic: Lacking, or living in the absence of, oxygen.
antagonism: Any type of interactive, interdependent relationship between
two or more organisms that is destructive to one of the participants.
antithesis, principle of: The observation that signals communicating op-
posite meaning tend to be expressed using displays having opposite
characteristics.
aposematic coloration: Bright warning coloration that toxic species use to
advertise their distastefulness to would-be predators.
aquatic ecology: The study of the ecology of freshwater systems (rivers,
lakes), estuaries, and marine environments (both coastal and open
ocean), including the physical, chemical, and biological processes asso-
ciated with them.
Arctic tundra: Treeless biome of very cold climates near to and north of the
Arctic Circle, in which the predominant plants are low-growing, peren-
nial woody plants and grasses. Lichens and mosses may also be com-
mon.
area-sensitive species: Species that require large blocks of natural area for
activities such as reproduction, finding food, and raising of young, are
called area-sensitive species. This required area may include portions of
open habitat, edge, and interior habitat. Area-sensitive species are espe-
cially sensitive to any reduction in area caused by habitat fragmentation.
688
Glossary
artificial selection: Choices made by plant breeders to produce varieties of

plants that have some desirable quality, such as improved yield, greater
height, or an unusual flower color.
asexual reproduction: Reproduction of cells or organisms without the
transfer or reassortment of genetic material, resulting in offspring that
are genetically identical to the parent.
autoecology: See physiological ecology.
autopolyploidy: Formation of a new species by the doubling of chromo-
somes of a single existing species. Many related species of plants within
a genus have been found to result from repeated occurrences of auto-
polyploidy.
autotrophs: Organisms that have the ability to make their own food from
inorganic substances. See also photosynthesis; primary producers.
back-cross: A cross involving an offspring individual crossed with one of

its parents.
banding: Technique for studying the movement, survival, and behavior of
birds by means of identification tags.
Batesian mimicry: An evolutionary trend in which an edible species mim-
ics the form of a distasteful species to avoid predation.
behavioral ecology: The systematic study of the strategies animals use to
overcome environmental problems and the adaptive value of those
strategies.
benthos: The area of the ocean floor; organisms associated with the sea
bottom.
bioaccumulation: See biomagnification.
bioclimatic zone: A zone of transition between differing yet adjacent eco-
logical systems.
biodegradable: Capable of being broken down, or degraded, into simpler
substances by natural decomposers.
biodiversity: This term represents an amalgamation of biology and diver-
sity. Ecologists typically recognize three types of biodiversity—species
biodiversity, genetic biodiversity, and habitat biodiversity. Habitat bio-
diversity refers to the variety of habitats in a given landscape, genetic
biodiversity refers to the number of alleles in a species genome or gene
pool in a given area. Species diversity refers to both the variety of spe-
cies and equal numbers of individuals among each species.
biogenetic law: Ernst Haeckel’s term for his generalization that the ontog-
eny of an organism recapitulates the adult stages of its ancestors (reca-
pitulation).
biogeochemical cycles: Movement of elements or water through both liv-
689
Glossary
ing and nonliving parts of an ecosystem. Carbon as carbon dioxide is

made into carbohydrate during photosynthesis and released through
decay to the nonliving atmosphere, from which it can later be reused in
photosynthesis.
biogeography: The science that seeks to understand spatial patterns of
biodiversity. See also island biogeography.
bioluminescence: Production of visible light by living organisms.
biomagnification: Also called bioaccumulation, the increasing accumula-
tion of a toxic substance in progressively higher feeding levels
biomass: The weight of organic matter in an environment or ecosystem, of-
ten expressed in terms of grams per square meter per year.
biomes: The primary, large-scale ecosystems of the world, largely identi-
fied with geographical regions typified by climate and weather and
classified in the Köppen system on the basis of precipitation, tempera-
ture, climate, soil types, flora, fauna, and location. Major biomes include
Tropical (rain forests, savannahs, tropical deciduous forests, and tropic
scrub), Mid-Latitude to Equatorial (temperate, hot, and cold grass-
lands, including steppe and chapparal; hot and cold deserts), Continen-
tal (woodlands, deciduous temperate forests, mediterranean woodland
and shrub), Moderate Continental (moderate grasslands, deciduous
forests, taiga and boreal forests), and Polar (ice caps, tundra).
biopesticides: Biological agents, such as viruses, bacteria, fungi, mites,
and other organisms used to control insect and weed pests in an envi-
ronmentally and ecologically friendly manner.
biosphere: Specifically, the 20-kilometer-thick zone extending from the
floor of the oceans to the top of mountains, within which all life on earth
exists. Generally, the sum of all the occupiable habitats for life on earth.
biota: All living things in a particular area, including microbes, fungi, al-
gae, plant life, and animals.
biotechnology: Combination of techniques whereby humans are able to
alter permanently the genetic makeup of organisms. Includes the in-
dustrial application of these techniques.
biotic: Living. Refers to the living components of an ecosystem or biome,
consisting of all organisms.
boreal forest: Located in two broad belts of vegetation that stretch from
east to west in the Northern Hemisphere, the primarily coniferous for-
ests that dominate this biome, which is also known as taiga.
bottleneck effect: In evolution, the reduction in size of a population caus-
ing a major loss of genetic variation. If the population size later ex-
pands, the new larger population will be genetically uniform and may
lack the ability to survive in a changing climate.
690
Glossary
brood: All the immature insects within an insect colony, including eggs,
larvae, and, in the Hymenoptera, the pupal stage; also, to cover young
with the wings.
brood parasite: See nest parasite.
browser: An animal that feeds on leaves and twigs from trees.
budding: A form of asexual reproduction that begins as an outpocketing of
the parental body, resulting in either separation from or continued con-
nection with the parent, forming a colony.
C3 plants: Plants whose system of photosynthesis produces a three-carbon

compound as the first identified compound after the uptake of carbon
dioxide during the light-independent reactions. See also CAM plants.
C4 plants: Plants whose system of photosynthesis produces a four-carbon
compound as the first identified compound after the uptake of carbon
dioxide during the light-independent reactions. C4 photosynthesis is
distinguished from CAM photosynthesis because C4 occurs during the
day and CAM occurs during the night. C4 plants are especially adapted
to hot, dry climates. Corn is an example. See also CAM plants.
calorie: The traditional unit of heat; one calorie is the amount of heat re-
quired to raise the temperature of one gram of water 1 degree Celsius.
CAM plants: Plants in desert biomes that use a crassulacean acid metabo-
lism to take in carbon dioxide during the night and store it as an acid,
and then use the carbon dioxide in the light-independent reactions dur-
ing the day, when sunlight is available. Cacti are CAM plants. See also C3
plants; C4 plants.
Cambrian explosion: The main period of evolutionary expansion in the
Phanerozoic era at the base of the Cambrian period, 544 million years
ago, which marks the development of all the modern phyla of organisms.
camouflage: Patterns, colors, and/or shapes that make it difficult to differ-
entiate an organism from its surroundings.
canopy: The uppermost portion of a rain forest, which shades the under-
story and the forest floor.
capture-recapture: See mark-capture-release method.
carbohydrates: Large class of organic molecules containing starch, carbon,
hydrogen, and oxygen and in which the ratio of hydrogen to oxygen is
two to one, the same as in a molecule of water. Sugars and cellulose are
examples.
carbon cycle: Biogeochemical cycle of the element carbon.
carbon fixation (CO2 ): Process by which carbon dioxide is made into glu-
cose during photosynthesis. This occurs during the part of photosyn-
thesis called the Calvin cycle.
691
Glossary
carcinogen: Any physical or chemical cancer-causing agent.

carnivore: A member of the meat-eating order Carnivora, which includes
dogs, cats, weasels, bears, and their relatives.
carnivorous plant: Plant that traps insects and digests them. These plants
usually live in nitrogen-poor habitats and use the insect proteins to sup-
plement their nitrogen intake.
carnivory: Subsisting or feeding on meat or flesh.
carrion: Dead animal flesh.
carrying capacity: The maximum number of animals that a given area can
support indefinitely.
caste: One of the recognizable types of individuals within an insect colony,
such as queens, workers, soldiers, and males or drones; usually these
individuals are physically and behaviorally adapted to perform specific
tasks.
catastrophism: A scientific theory which postulates that the geological fea-
tures of the earth and life thereon have been drastically affected by nat-
ural disasters of huge proportions in past ages.
cellulose: A fibrous polysaccharide that chiefly constitutes the cell walls of
plants and is not easily dissolved in water. Primary consumers, includ-
ing ruminants, require specialized digestive systems to break down cel-
lulose.
census: The counting of populations of naturally occurring organisms to
understand their ecology more fully.
CFCs: See Chlorofluorocarbons (CFCs).
chaparral: Biome found along the coast of Southern California, char-
acterized by short trees with leathery leaves, shrubs, and open grassy
areas.
character displacement: A change in the morphological, behavioral, or
physiological state of a species without geographical isolation, as a re-
sult of natural selection arising from competition between one or more
ecologically similar species.
chemical ecology: Ecology that concerns the biochemicals (called semio-
chemicals) produced and released by organisms that have physiologi-
cal and behavioral effects on other organisms. Studies in chemical ecol-
ogy are often interdisciplinary (integrating several fields of science such
as chemistry and ecology) but also more specific areas in biochemistry
(such as biosynthesis of compounds), molecular biology, or physiology
(reception of the compounds and transmission of nerve impulses), as
well as in behavioral ecology (orientation movements of an organism)
and population ecology (aggregation and competition of organisms)
and even the interactions among trophic levels (such as predator-prey
692
Glossary
interactions). Evolutionary studies at all these levels are of interest to

understand how stable the semiochemical systems are and whether ad-
aptations to new systems are constrained.
chemical pollutants: Harmful chemicals manufactured and released to the
environment.
chemosynthetic autotrophs: Organisms (usually bacteria) that make com-
plex food molecules from simpler molecules using energy of chem-
ical reactions rather than light energy used by photosynthetic auto-
trophs.
chlorofluorocarbons (CFCs): A group of very stable compounds used
widely since their development in 1928 for refrigeration, coolants, aero-
sol spray propellants, and other uses; once risen in stratosphere, they
cause ozone depletion.
circadian rhythm: A physiological or behavioral cycle that occurs roughly
in a twenty-four-hour pattern.
cladistics: System of describing evolutionary relationships in which only
two groups, or clades, branch from each ancestral group. The more re-
cently two clades diverged, the more characteristics they have in com-
mon and the closer they will appear in a cladistic diagram.
class: The taxonomic category composed of related genera; closely related
classes form a phylum or division.
classification: The arranging of organisms into related groups based on
specific relationships. See also systematics; taxonomy.
clear-cutting: The removal of all trees from an area.
climax community: Group of plants that appear late in succession and are
not replaced by plants of different species unless the climate changes or
the area is disturbed, as by fire or cultivation.
cline: A graduated series of populations of the same species. Each popula-
tion has a slightly different physiology from the ones on each side.
Clines typically develop where environmental factors change in a grad-
ual way, such as from the bottom of a mountain to the top.
clone: An organism that is genetically identical to the original organism
from which it was derived.
cloning: The technique of making a perfect genetic copy of a DNA mole-
cule, a cell, or an entire organism.
clutch: A group of eggs laid in a single reproductive effort.
coevolution: Simultaneous evolutionary change through time of two spe-
cies, such as a flower and its pollinating insect, each influenced by the
changes the other species is undergoing. Over a period of time, the two
species often become dependent on each other, so that one would not
survive the disappearance of the other.
693
Glossary
cognitive ethology: The study of animal intelligence.

cohort: A group of organisms of the same species, and usually of the same
population, that are born at about the same time.
colony: A cluster of genetically identical individuals formed asexually
from a single individual.
coloration: See aposematic coloration; cryptic coloration.
commensalism: A type of coevolved symbiotic relationship between dif-
ferent species that live intimately with one another without injury to
any participant.
communication: The exchange of information between members of a spe-
cies by means of chemical signals (pheromones), displays, calls, and
other means.
community: A population of plants and animals that live together and in-
teract with one another through the processes of competition, preda-
tion, parasitism, and mutualism, making up the biotic part of an ecosys-
tem.
community ecology: The study of the impacts that populations of different
species have on populations of other species with which they interact,
be those interactions between plants and other plants, animals and
other animals, or plants and animals. The emphasis is on how these
populations of different species change, enhance, or delimit one an-
other. Population ecology is related but is focused on the growth and
change in populations of discrete species.
comparative physiology: See physiological ecology.
compartmentalization: A characteristic of most communities, in which a
given set of producers tends to be consumed by a limited number of
consumers, which in turn are preyed upon by a smaller number of pred-
ators, and so on.
competition: The interactions among individuals that attempt to utilize
the same limited resource.
competitive exclusion, principle of (Gause’s principle): If two or more
species compete for the same niche, one of them will be successful, and
the other will be eliminated over time.
coniferous forest: Large group of trees that are predominantly conifers,
such as a pine forest or a spruce-fir forest.
conjugation: Type of sexual reproduction that occurs in green algae such
as Spirogyra and in certain kinds of fungi. Also, the transfer of ge-
netic material from one bacterium to another through a cytoplasmic
bridge.
connectivity: The ability of organisms to use corridors of habitat to dis-
perse from one habitat patch to another.
694
Glossary
conservation biology: The use of biological science to design and imple-

ment methods to ensure the survival of species, ecosystems, and ecologi-
cal processes. Conservation biologists are concerned with the process of
speciation, the measurement of biodiversity, and factors involved in the
extinction process. However, the primary thrust of their efforts is the
development of strategies to preserve biodiversity; hence, conservation
biology is a value-laden science.
conservation easement: An arrangement in which a national government,
private organization, or consortium of countries compensates a tropi-
cal country for protecting a specific habitat. See also debt-for-nature
swap.
consort pair: A temporarily bonded pair within a polygamous group; also
called consortship.
conspecific: A member of the same species.
constriction: A method of killing prey using increasingly tight coils
around the body to trigger stress-induced cardiac arrest.
consumer: An organism other than a primary producer—that is, one that
eats other organisms, including primary consumers (herbivores), sec-
ondary consumers (omnivores and carnivores), scavengers, and decom-
posers.
continuous growth: Growth in a population in which reproduction takes
place at any time during the year rather than during specific time inter-
vals.
convergent evolution: The process by which evolutionarily unrelated ani-
mals tend to resemble one another as a result of adaptations to similar
environments.
cooperation: A social behavior in which members of a group act for the
good of all.
coral reef: A reef built primarily by coral species.
core species: A species that utilizes the interior area or core of a habitat.
Core species such as many neotropical migrants typically require large
blocks of contiguous natural habitat for reproduction and are typically
very sensitive to the ratio of core-to-edge habitat in a given patch or par-
cel of landscape.
corridor: Narrow link of habitat that connects two or more patches in a
landscape. Natural corridors serve as wildlife dispersal conduits, per-
mitting movement between natural habitats with minimal risk of expo-
sure.
countershading: A form of crypsis involving dark coloration on top and
light coloration on the underside.
coupled oscillations: In predator-prey relationships, the waxing and wan-
695
Glossary
ing of population sizes of two species in a community, based on cycles

of predation.
crassulacean acid metabolism (CAM): See CAM plants.
crepuscular: Active after sunset and in early morning.
Cretaceous-Tertiary (KT) event: An event that occurred about 66.4 million
years ago, sometimes hypothesized to be a meteoritic impact, that marks
the boundary between the Cretaceous and Tertiary periods and that ini-
tiated the mass extinctions of many species, notably the dinosaurs.
critical period: A very brief period of time in the development of an animal
during which certain experiences must be undergone; the effects of
such experiences are permanent.
critical photoperiod: Specific day length necessary to produce flowers in
long-day and short-day plants.
cross-pollination: The transfer of pollen grains and their enclosed sperm
cells from the male portion of a flower to a female portion of another
flower within the same species.
crown: The branched, leafy part of a tree.
crypsis: The phenomenon of hiding or remaining hidden.
cryptic coloration: Any color pattern that blends into the background.
cuckold: A partnered male who is helping his mate to raise offspring
which are not genetically his own.
cud: Food regurgitated and chewed a second time after its initial ingestion.
cultural diversity: Variety of learned behaviors among individuals of a
species.
cyanobacteria: Photosynthetic prokaryotes that were once called blue-
green algae.
debt-for-nature swap: An arrangement in which tropical countries act as

custodians of the tropical forest in exchange for foreign aid or relief
from debt. See also conservation easement.
deciduous tropical forest: Forests of tropical regions that shed their leaves
during annual dry periods.
decomposers: Bacteria and fungi that break down dead organic matter. In
the process, they obtain energy and facilitate the recycling of elements.
deep ecology: A philosophy, introduced by the Norwegian philosopher
Arne Naess, that values the natural world and biodiversity in and of
themselves and advocates seeing nature as more than a resource for hu-
man use.
defense mechanism: Any of a variety of chemical, behavioral, anatomical,
or physiological means by which an organism prevents or discourages
predation.
696
Glossary
definitive host: The host in which a symbiont (the organism living within
the host) matures and reproduces.
defoliant: A chemical that kills the leaves of trees.
deforestation: The removal of trees from forests to an extent that degrades
the forest biome without human intervention. See also reforestation.
deme: A local population of closely related living organisms.
demography: The study of the numbers of organisms born in a population
within a certain time period, the rate at which they survive to various
ages, and the number of offspring that they produce. Also referred to as
demographics.
dendrochronology: The examination and comparison of growth rings in
both living and aged woods to draw inferences about past ecosystems
and environmental conditions.
dendroclimatology: The study of tree-ring growth as an indicator of past
climates.
denitrification: Process in which bacteria convert nitrogenous compounds
in the soil to nitrogen gas.
denning: The period of winter sleep during which a bear does not eat,
drink, urinate, or defecate.
density: The number of animals present per unit of area being sampled;
for example, ten mice per hectare or five moose per square kilometer.
density-dependent growth: Growth in a population in which the per ca-
pita rates of birth and death are scaled by the total number of individu-
als in the population.
density-dependent population regulation: The regulation of population
size by factors or interactions intrinsic to the population; the strength of
regulation increases as population size increases.
deoxyribonucleic acid (DNA): The genetic material of cells, having the
molecular form of a twisted double helix that is linked by purine and
pyrimidine base pairs; carries the inherited traits and controls for cell
activities.
desert: Biome that receives less than 10 inches of precipitation per year.
desertification: The degradation of arid, semiarid, and dry, subhumid
lands as a result of human activities or climatic variations, such as a pro-
longed drought.
despotism: A type of hierarchy in which one individual rules over all other
members of the group and no rank distinctions are made among the
subordinates.
detritus feeders (detritivores): An array of small and often unnoticed ani-
mals and protists that live off the refuse of other living beings, such as
molted shells and skeletons, fallen leaves, wastes, and dead bodies.
697
Glossary
diapause: A resting phase in which metabolic activity is low and adverse

conditions can be tolerated; also, an interruption in embryonic develop-
ment.
diatom: Microscopic algae that produce a frustule (a kind of shell) made of
silica glass that is highly resistant to weathering. A major type of
phytoplankton.
differentiation: The process during development by which cells obtain
their unique structure and function.
digestion: The process by which larger organic nutrients are broken down
to smaller molecules in the lumen of the gut.
dilution effects: The reduction in per capita probability of death from a
predator due to the presence of other group members.
dimorphism: Existence of two distinct forms within a species.
dinoflagellates: A unicellular, mobile type of phytoplankton responsible
for deadly algal blooms called red tides.
dioecious: Having two separate sexes, namely male and female.
diploid: Having two sets of chromosomes, usually one derived from the
father and one derived from the mother; the normal condition of all
cells except reproductive ones. See also haploid.
discrete growth: Growth in a population that undergoes reproduction at
specific time intervals.
discrete signals: Signals that are always given in the same way and indi-
cate only the presence or absence of a particular condition or state.
disease ecology: Ecological factors influencing, and influenced by, the
emergence and spread of infectious diseases, including both plant and
animal populations. Considers such questions as how biodiversity af-
fects the spread of disease, patterns of disease spread through popula-
tions, and the roles of evolution and genetics.
disharmonic: Ecologically unbalanced.
disjunct: Pertaining to the geographic distribution pattern in which two
closely related groups are widely separated by areas that are devoid of
either group.
dispersal: The movement of organisms from one geographic area to an-
other; movements may be the result of an animal’s own efforts (active
dispersal) or the consequence of being transported by natural or hu-
man-mediated means (passive dispersal) and can be limited by physi-
cal barriers.
dispersion: The pattern or arrangement of members of a population in a
habitat; also, the transport or movement of seeds across a substrate such
as soil prior to germination.
698
Glossary
disphotic marine zone: A transitional marine region between the photic

and aphotic zones that may extend to depths of one thousand meters;
also called the mesopelagic zone.
display: A social signal, particularly a visual signal, exchanged between
animals.
disruptive coloration: Use of stripes, spots, or blotches to break up the
body outline and blend into a complex background.
diurnal: Awake and functional during the daylight hours.
divergence: The evolution of increasing morphological differences be-
tween an ancestral species and offshoot species caused by differing
adaptive pressures.
diversity: The number of taxa (classification groups) associated with a par-
ticular place and time. See also biodiversity.
DNA: See deoxyribonucleic acid.
domestication: A process by which animals are adapted biologically and
behaviorally to a domestic (human) environment in order to tame and
manipulate them for the benefit of humans.
dominance: The physical control of some members of a group by other
members, initiated and sustained by hostile behavior of a direct, subtle,
or indirect nature.
dominance hierarchy: A social system, usually determined by aggressive
interactions, in which individuals can be ranked in terms of their access
to resources or mates.
dominant species: A species in a community that acts to control the abun-
dance of its competitors because of its large size, extended life span, or
ability to acquire and hold resources.
dormancy: A period of inactivity that allows a plant or animal to survive
unfavorable cold or dryness (hypernation, estivation).
drone: A fertile male social insect.
drumming: Type of nonvocal communication that a bird produces by
banging its bill on a hollow tree trunk or other noise-producing object.
dry tropical biomes: The savanna and deciduous tropic forest biomes,
which often occur where the annual rainfall is less than that of savan-
nas, most often found in Africa, South America, and Australia.
dynamic equilibrium: Characterizing a community in equilibrium that is
always responding to the last disturbance.
dynamically fragile: Characterizing a community that exhibits resilience
only within a narrow range of conditions.
dynamically robust: Characterizing a communitiy that exhibits resilience
over a wide range of conditions and scales of disturbance.
699
Glossary
eclipse plumage: The drab plumage of male birds following the post-
breeding molt, in which their bright courtship feathers are replaced by
dull earthy feathers that provide inconspicuous coloring.
ecocentrism: A philosophy that emphasizes the value of nature as a whole
and an identification of the self with the natural world.
ecoenergetics: The flow of energy through ecological systems at all levels,
from individual organisms, populations, and communities to ecosys-
tems and the global environment. Includes abiotic factors (such as geo-
chemical cycles) as well as biotic factors.
ecofeminism: A consideration of gender differences in the experience of
the self and nature, which includes an analysis of the tie between the op-
pression of women and nature.
ecological niche: See niche.
ecology: The study of the interactions between organisms and the living
(biotic) and nonliving (abiotic) components of their environment, in-
cluding the distribution and abundance of organisms.
ecomorph: Species of different phyletic origins (at most distantly related)
with similar structural and behavioral adaptations to similar niches.
ecophysiology: See physiological ecology.
ecosystem: A biological community and the physical environment con-
tained in it.
ecosystem diversity: Variety of biomes and habitats occuring in the bio-
sphere.
ecosystem ecology: The study of the flow of energy into, through, and out
of large-scale systems, and how that flow influences all abiotic factors
and living organisms in the ecosystem.
ecotone: The meeting place between the two biomes, a transition zone.
ecotourism: Tourism based on the promotion of ecologically important
sites, such as national parks, wildlife refuges, endangered ecosystems,
and the organisms for which they form the habitat.
ecotoxicology: The study of natural and man-made pollutants and their
toxic effects on organisms, populations, communities, and ecosystems,
as well as the ways these pollutants impact ecological processes to
change ecosystems and their components.
ectoparasite: A parasitic organism that attaches to the host on the exterior
of the body.
edge species: Certain species of wildlife that exploit edge habitat, which is
the transition habitat zone between two distinct habitats.
elfin forest: A stunted forest growing at high elevations in warm, moist cli-
mates.
emergent vegetation: Vegetation that grows tall enough to be visible at
700
Glossary
the highest level of a rain forest or, in aquatic systems, above the water
line.
emigration: The movement of animals out of an area; one-way movement
from a habitat type.
encounter effects: The reduction in the probability of death from a preda-
tor due to a single group of N members being more difficult to locate
than an equal number of solitary individuals.
endangered species: A species of animal or plant, as designated by bodies
such as the U.S. Fish and Wildlife Service, that is threatened with extinc-
tion.
endemic: Belonging to or native to a particular place; often referring to
species that have evolved in a given area and are found nowhere else in
the world.
endosymbiosis: A symbiotic relationship in which one member lives in-
side the other’s body.
endocannibalism: A form of human cannibalism in which members of a
related group eat their own dead.
endoparasite: A parasitic organism that attaches to an interior portion of
the host’s body.
endophyte: An organism living within a plant, such as a fungus living in a
root.
endosymbiotic theory: Theory that chloroplasts and mitochondria devel-
oped from bacteria that moved into and became essential to the survival
of early eukaryotic cells.
endotherm: An animal that, by its own metabolism, maintains a constant
body temperature (warm-blooded); birds and mammals are endo-
therms.
energy: The ability to do work. Energy takes several forms, such as chemi-
cal energy in the bonds of a compound, light energy, and kinetic energy,
the energy of movement.
energy budget: The amount of resources available to an organism, which
is accepted to be limited or finite. Energy acquired from food (ani-
mals) or sunlight (plants) must be partitioned among growth, main-
tenance, and reproduction. The greater the energy allocated to the
care of offspring, for example, the fewer the offspring that can be pro-
duced.
energy flow: The capture of radiant energy, its transformation into chemical
energy by producers via photosynthesis, and its translocation through
all biological systems via consumers and decomposers. All organisms
are considered as potential sources of energy. See also food chain.
energy pyramid: A graphical representation of the energy contained in
701
Glossary
succeeding trophic levels, with maximum energy at the base (produc-

ers) and steadily diminishing amounts at higher levels.
entrainment: The synchronization of one biological rhythm to another
rhythm, such as the twenty-four-hour rhythm of a light-dark cycle.
entropy: The tendency of complex molecules and other structures to lose
energy and become degraded into simpler forms.
environment: All the external conditions that affect an organism or other
specified system during its lifetime.
environmental constraints: The physical demands placed upon any spe-
cies by its surroundings that ultimately determine the success or failure
of its adaptations and consequently its success as a species; also called
pressures.
epifauna: Animals that live on the sea floor.
epiphyte: A plant that lives nonparasitically on another plant. Usually,
epiphytes are not rooted in the ground and are typically found in habi-
tats with high humidity. Spanish moss is an epiphyte.
epizoites: Commensals that live on the skin, fur, scales, or feathers of their
hosts.
estivation: Similar to hibernation, a period of reduced activity or dor-
mancy triggered by dry or hot environmental conditions. See also hiber-
nation.
estuarine ecosystem: An aquatic ecosystem that occurs where a freshwa-
ter river meets a saltwater or ocean environment.
ethology: The study of an animal’s behavior in its natural habitat.
euphotic zone: Also called the photic zone, the region of a body of water
that is penetrated by sunlight.
eusocial: Characterizing a social system with a single breeding female;
other members of the colony are organized into specialized classes (ex-
emplified by bees, ants, and termites).
eutrophication: The overenrichment of water by nutrients, causing exces-
sive plant growth and stagnation, which in turn leads to the death of
fish and other aquatic life.
evapotranspiration: The loss of water by means of both evaporation from
soil and transpiration from plants.
evergreen: Plant that keeps its leaves during adverse climatic conditions,
such as cold temperatures. Examples are pine trees and rhododendrons.
evolution: A process, guided by natural selection, that changes a popula-
tion’s genetic composition and results in adaptations.
evolutionarily stable strategy: A behavioral strategy that will persist in a
population because alternative strategies, in the context of that popula-
tion, will be less successful.
702
Glossary
evolutionary ecology: The study of how evolutionary processes such as

selection and adaptation influence the interactions of organisms with
their environments and shape species and ecosystems.
exogenous: Originating outside an ecosystem, community, population or
organism.
exotic species: Organisms that are not naturally found in a place but have
been artificially introduced, whether by accident or intentionally. See
also invasive species.
exponential growth: A pattern of population growth in which the rate of
increase becomes progressively larger over time.
extant: Alive and reproducing, as opposed to extinct species.
external fertilization: The union of eggs and sperm in the environment,
rather than in the female’s body.
extinct: No longer living on earth.
F1 generation: First filial generation; offspring produced from a mating of

P generation individuals.
F2 generation: Second filial generation; offspring produced from a mating
of F1 generation individuals.
facultative anaerobe: Organism that can survive in an oxygen-poor envi-
ronment when necessary, using the small amount of energy available
from fermentation.
fecundity: The number of offspring produced by an individual.
female: An organism that produces eggs, the larger of two different types
of gametes.
fermentation: Set of reactions that change glucose into alcohol and carbon
dioxide. A small amount of energy is produced, some of which is cap-
tured as ATP.
fertilization (agriculture): Addition of minerals or decayed organic mat-
ter to soil that has been depleted of nutrients by farming or other
means.
fertilization (sexual reproduction): The fusion of sperm and egg to form a
fertilized egg (zygote).
field capacity: Water remaining in soil following rain or irrigation after ex-
cess water has been drained off by gravity.
fire climax ecosystem: An ecosystem that depends on periodic fires to
clear underbrush; the seeds of many plants in such an ecosystem re-
quire fire in order to germinate.
fire ecology: The study of how both natural and planned fires impact eco-
systems such as forests.
fitness: The ability of an organism to produce offspring that, in turn, can
703
Glossary
reproduce successfully; the fitness of organisms increases as a result of

natural selection.
fixation: Process by which a gas has been converted to another type of
molecule, usually organic.
fledgling period: Period after hatching, during which a nestling grows
flight feathers and learns to fly.
food chain: A paradigm representing the links between organisms, each of
which eats and is eaten by another.
food pyramid: A diagram representing organisms of a particular type that
can be supported at each trophic level from a given input of solar en-
ergy in food chains and food webs.
food web: A network of interconnecting food chains representing the food
relationships in a community.
foraging: See active or wide foraging; sit-and-wait foraging.
fossil: A remnant, impression, or trace of an animal or plant of a past geo-
logical age that has been preserved in the earth’s crust.
fossil fuel: A fuel product originating from the partial or complete decom-
position of carbon-based life-forms (plant and animal remains and fos-
sils) exposed to heat and pressure in the earth’s crust over thousands
and millions of years. Fossil fuels include crude oil, coal, natural gas,
and gasoline.
fossil record: The evolutionary information contained in the fossils found
in the earth’s crust when compared with the geologic record.
frequency-dependent predation: Predation on whichever species is most
common in a community; a frequency-dependent predator will switch
prey if necessary.
frozen zoo: A frozen tissue bank, maintained at many zoos, that contains
wild animal tissue and reproductive samples for use in future breeding
programs.
functional response: The rate at which an individual predator consumes
prey, dependent upon the abundance of that prey in a habitat.
fusion-fission community: A society whose members are of both sexes
and all ages, which can form and dissolve subgroupings.
gamete: A haploid reproductive cell, usually a sperm or egg.

Gause’s principle: See competitive exclusion, principle of.
gene: A portion of a DNA molecule containing the genetic information
necessary to produce a molecule of messenger RNA (via the process of
transcription) that can then be used to produce a protein (via the pro-
cess of translation).
gene flow: The movement of genes from one population to another through
704
Glossary
migration and hybridization between individuals belonging to adjacent

populations.
gene frequency: The occurrence of a particular allele present in a popula-
tion, expressed as a percentage of the total number of alleles present.
gene (point) mutation: A change within the hereditary material of a single
gene. These tiny mutations cannot be observed by inspecting pictures of
chromosomes.
gene pool: All the alleles of all the genes present in a population. There is
no limit to the number of alleles in a gene pool; however, an individual
may not possess more than two different alleles.
generalized: Not specifically adapted to any given environment.
genetic diversity: The total number and distribution of alleles and geno-
types in a population; a population with a very high genetic diversity
would have many alleles and genotypes, all evenly distributed or with
approximately equal frequency.
genetic drift: Change in gene frequencies in a population owing to chance.
genetic modification: Alteration of a organism’s genetic material by ma-
nipulation in the laboratory. The addition of new genes or the removal
of genes are examples.
genome: The complete amount of DNA found in the nucleus of a normal
cell, expressed as a particular number of chromosomes; for example, a
human cell has a genome of forty-six chromosomes.
genotype: The complete genetic makeup of an organism, regardless of
whether these genes are expressed. See also phenotype.
genotype frequency: The relative abundance of a genotype in a popula-
tion; to calculate, count the number of individuals with a given geno-
type in the population and divide by the total number of individuals in
the population.
genus (pl. genera): A group of closely related species having many traits in
common and descended from a common ancestor; for example, Felis is
the genus of cats, and it includes the species Felis catus (the domestic cat)
and Felis couguar (the cougar or mountain lion).
geochemical cycles: See biogeochemical cycles.
global ecology: The study of the impacts of such factors as global warm-
ing, pollution, and disease on organisms and ecosystems worldwide.
Much of global ecology considers the ecological impacts of human-
driven influences such as international travel, trade, the built environ-
ment, and the use of petrochemicals.
global extinction: The loss of all members of a species; that is, extinction
whereby all populations of a species disappear or are eliminated. See
also local extinction.
705
Glossary
global warming: The theory that the atmosphere is becoming warmer over
time as a result of an increase in greenhouse gases, resulting in climate
change, melting ice caps, rising sea levels, severe weather events, and
their impacts on the world’s living organisms.
glycosides: Compounds produced by plants that combine a sugar, usually
glucose, with an active component. Potentially poisonous glycosides
include the cyanogenic, cardioactive, anthraquinone, coumarin, and
saponin glycosides.
gradient analysis: A method for studying the distribution of species along
an environmental gradient.
gradualism: A model of evolution in which transformation from ancestor to
descendant species is a slow, gradual process spanning millions of years.
grassland: A biome in which the dominant plants are grasses.
grazer: An organism that feeds primarily on grasses.
Green Revolution: Several decades of dramatic advances in yield and
quality of crop species. This was the outcome of attempts to increase
food production begun in the 1940’s.
greenhouse effect: The process whereby the infrared radiation from the
earth is absorbed by the atmosphere, keeping it from escaping into
space. Because it was once believed that the glass panes of greenhouses
acted similarly, this phenomenon was termed the “greenhouse” effect,
although it has subsequently been shown that greenhouses work differ-
ently (by trapping heated air and not allowing it to blow away).
greenhouse gases: gases, including carbon dioxide, water vapor, and
methane, store heat more efficiently than others and contribute to the
magnification of the so-called greenhouse effect.
gregarious: Forming groups temporarily or permanently.
gross primary productivity: The amount of the sun’s energy actually as-
similated by autotrophs. See also net primary productivity; secondary
productivity.
habitat: The physical environment, usually that of soil and vegetation as

well as space, in which an animal lives.
habitat fragmentation: Conversion of a natural, contiguous landscape into
smaller patches usually due to human activity. Grids of roads, gas lines,
cluster housing, and power lines all contribute to habitat fragmentation.
habitat selection: Process of choosing a home range, territory, nesting site,
or feeding site on the basis of specific features of the habitat that an or-
ganism is best adapted to exploit.
habituation: The process of learning to ignore irrelevant stimuli that previ-
ously produced a reaction.
706
Glossary
haploid: Having one set of chromosomes and one of each kind of gene. Ga-
metes (eggs or sperm) are usually haploid. See also diploid.
Hardy-Weinberg law: A concept in population genetics stating that, given
an infinitely large population that experiences random mating without
mutation or any other such affecting factor, the frequency of particular
alleles will reach a state of equilibrium, after which their frequency will
not change from one generation to the next.
hazardous waste: Any waste product that is toxic to living beings or
threatens life.
hemiparasite: A plant (such as mistletoe) with some chlorophyll which
lives as a partial parasite.
herbicide: Chemical that is lethal to plants.
herbivore: An animal that eats living plants, usually specialized to digest
cellulose.
heritability: The extent to which variation in some trait among individuals
in a population is a result of genetic differences.
heterochrony: Any phenomenon in which there is a difference between
the ancestral and descendant rate or timing of development.
heteroecious: Describes fungi that spend parts of their life cycle on entirely
different species of plants. Black stem rust of wheat spends part of its
life cycle on wheat and part on American barberry.
heterotrophs: Organisms that cannot make their own food from simpler
materials but must ingest and digest complex molecules made by other
organisms. Animals are heterotrophs, but so are nongreen plants such
as Indian pipes. See also consumer.
heterozygous: Having two different alleles at a particular gene locus.
hibernation: A sustained period of torpor (lack of activity) triggered by
cold environmental conditions, achieved when an animal reduces its
metabolic rate. See also estivation.
hierarchy: A social structure in which animals are dominated by those
higher on the linear ladder.
home range: Geographic area used by an individual, pair, or group for
their daily, seasonal, and sometimes their yearly activities; the defended
portion of the home range is called a territory.
homeostasis: The dynamic balance between body functions, needs, and
environmental factors which results in internal constancy.
homologous: Referring to chromosomes that are identical in terms of
types of genes present and the location of the centromere; because of
their high degree of similarity, homologous chromosomes can synapse
and recombine during prophase I of meiosis.
homozygote: A diploid organism that has two identical alleles for a partic-
707
Glossary
ular trait; a person with blood type A would be homozygous if he had

two A alleles.
horizons (soil): Layers of soil with different appearance and different
chemical composition. In a forest, the horizon closest to the top will con-
tain more organic matter than will the deeper layers.
hormone: Chemical compound produced in small amounts in one part
of an organism that has an effect in a different part of the same organ-
ism. Auxins, gibberellins, cytokinins, ethylene, estrogen, progesterone,
oxytocin, and various pheromones are examples.
host: In a parasitic relationship, the organism that is giving up energy-
containing molecules to the parasite.
hybrid: An organism resulting from the crossing of two species.
hybrid vigor: The tendency of hybrids to be larger and more durable than
their parent species; also called heterosis.
hybrid zone: An area with a population of a species composed of individu-
als with characteristics of one or more species that have interbred.
hybridization: A process of base-pairing involving two single-stranded nu-
cleic acid molecules with complementary sequences; the extent to which
two unrelated nucleic acid molecules will hybridize is often used as a
way to determine the amount of similarity between the sequences of the
two molecules; hybridization is fairly common among wind-pollinated
plants, while hybridization is quite uncommon among higher animals.
hydrologic cycle: Earth’s cycle of evaporation and condensation of water,
which produces rain and maintains oceans, rivers, and lakes.
hydrophilic: Loving water, capable of mixing with or growing in water.
hydrosphere: The waters of the earth and the regions where they are
found, including freshwater lakes and rivers, the oceans, inland seas,
and then frozen water of the polar zones.
immigration: The movement of organisms into an area; a one-way move-

ment into a habitat type.
imprinting: A specialized form of learning characterized by a sensitive pe-
riod in which an association with an object is formed.
inbreeding: Mating between relatives, an extreme form of positive assor-
tative mating. See also outbreeding.
inbreeding depression: Weakening, or lowering of the fitness, of offspring
as a result of inbreeding, caused when alleles that decrease fitness drift
to fixation.
indicator species: Any species that is among the first to be degraded when
an ecosystem is compromised, indicating the ecological impact of envi-
ronmental change.
708
Glossary
individual ecology: See behavioral ecology; physiological ecology.

industrial melanism: The rapid rise in frequency of the melanic form in
many moth species downwind of manufacturing sites, associated with
the advent of industrial pollution.
infauna: Animals that live in the sea floor.
inflorescence: The group of flowers that forms at the top of a flower stalk.
Inflorescences may be compact, like that of a daisy, or loose, like that of a
mustard.
innate: Inborn, possessed from birth, or determined and controlled largely
by the genes.
insectivore: Any of an order of small, nocturnal mammals, including
shrews, moles, and hedgehogs, Insectivora, or generally any animal that
feeds on insects.
insectivorous plant: Plant that traps insects and digests them for the nitro-
gen-containing compounds they possess. Plants of this type grow in
boggy soil that is typically low in nitrogen.
instinct: Any behavior that is completely functional the first time it is per-
formed.
integrated pest management (IPM): The practice of integrating insect, ani-
mal, or plant management tactics, such as chemical control, cultural
control, biological control, and plant resistance, to maintain pest popu-
lations below damaging levels in the most economical and environmen-
tally responsible manner.
interbreeding: The mating of closely related individuals which tends to in-
crease the appearance of recessive genes.
interference: The act of impeding others from using some limited resource.
interfertile: Able to breed and produce fertile offspring.
intermediate host: An animal species in which nonsexual developmental
stages of some commensals and parasites occur.
interspecific competition: Competition between species that need the
same limited resource.
intertidal marine zone: The shallow marine biome that lies at the land’s
edge.
intraspecific competition: Competition between members of the same
species for a limited resource.
intrinsic rate of increase: The growth rate of a population under ideal con-
ditions, expressed on a per individual basis.
introgression: The assimilation of the genes of one species into the gene
pool of another by successful hybridization.
invasive species: Nonnative (also termed “exotic”) organisms that can
outcompete native species. See also exotic species.
709
Glossary
iridescent: Showing the colors of the rainbow depending light reflection.

irruption: A sudden increase in the size of a population, usually attributed
to a particularly favorable set of environmental conditions.
island biogeography: a distinct subdiscipline of biogeography that con-
siders biodiversity and island size, ecological heterogeneity, proximity
to continents, isolation and endemism, island size and location and
rates of immigration, colonization, and extinction. Island biogeogra-
phers frequently have debates arguing the relevance of dispersal versus
vicariance.
isolation: See reproductive isolation.
isotherm: A line or boundary imagined on the earth’s surface that connects
points having the same temperature at a given time.
K strategy [selection]: A reproductive strategy typified by low reproduc-

tive output; common in species living in areas having limited critical re-
sources.
keystone species: A species that determines the structure of a commu-
nity, usually by predation on the dominant competitor in the commu-
nity.
kin selection: A phenomenon by which acts of altruism can help pass
on genes for altruism by improving the survival of kin and their off-
spring.
landscape ecology: A relatively new field of ecology, the study of how eco-
systems, including the built environment, are arranged and how their
arrangements affect the wildlife and environmental conditions that
form them. Landscape ecologists examine land patterns (topography,
water, forest cover, and human uses) and how these affect wildlife pop-
ulations.
lichen: An organism formed by the symbiotic relationship between a fun-
gus and an alga. The alga is protected from drying by the fungus, while
the fungus receives food molecules from the alga.
life cycle: The sequence of development beginning with a certain event in
a organism’s life (such as the fertilization of a gamete), and ending with
the same event in the next generation.
life expectancy: The probable length of life remaining to an organism
based upon the average life span of the population to which it belongs.
life span: The maximum time between birth and death for the members of
a species as a whole.
life table: A chart that summarizes the survivorship and reproduction of a
cohort throughout its life span.
710
Glossary
limnology: The study of the physical, chemical, climatological, biological,

and ecological aspects of lakes.
litter: The offspring produced in a single birth; also referred to as a clutch.
littoral zone: The shore zone of a lake, where macrovegetation grows.
local extinction: The loss of one or more populations of a species, but with
at least one population of the species remaining. See also global extinc-
tion.
logistic growth: A pattern of population growth that involves a rapid in-
crease in numbers when the density is low but slows as the density ap-
proaches the carrying capacity.
macroevolution: Large-scale evolutionary processes that result in major

changes in organisms and allow them to occupy new adaptive niches or
develop novel body plans.
marine ecology: A branch of both ecology and oceanography that investi-
gates the ocean zones and the interactions of their biotic and abiotic
components, including all components impinging on the oceans such as
seabirds and coastal zones.
mark-capture-release methods: Methods of studying populations by cap-
turing and marking some members, then releasing them into the wild
and periodically recapturing them, or capturing successive samples, in
order to track their status at different points in time.
mass extinction: An event in which a large number of organisms in many
different taxa are eliminated; there have been five such events in the his-
tory of life that resulted in the disappearance of more than 75 percent of
all species.
mate competition: Competition among members of one sex for mating op-
portunities with members of the opposite sex.
maximum sustainable yield (MSY): The rate of harvest of natural re-
sources such as fisheries and timber that can be maintained indefinitely
through active human management of those resources.
mediterranean scrub: Type of vegetation found in certain places, such as
Southern California, which experience wet winters and long, dry sum-
mers.
metabolites: Compounds formed as the result of biochemical pathways in
an organism.
metamorphosis: An abrupt change from one life-form to another, such as
from a larval body form, accompanied by many physiological changes
in the determination, differentiation, and distribution of cells, into an
adult body form.
microbivores: Organisms that eat microbes.
711
Glossary
microevolution: Small-scale evolutionary processes resulting from grad-

ual substitution of genes and resulting in very subtle changes in organ-
isms.
microphages: Animals that feed on small microscopic particles suspended
in water or deposited on bottom sediments.
migration: The movement of individuals resulting in gene flow, changing
the proportions of genotypes in a population.
mimicry: The resemblance of one species (the model) by one or more
other species (mimics), such that a predator cannot distinguish among
them.
molecular clock: Accumulation of genetic changes that develops when
two species diverge. The longer two species have been separate, the
more changes will be evident. The clock does not tick at the same rate in
every species or every molecule.
molecular ecology: The study of natural and introduced microbial popula-
tions and their environments and ecological implications of the release
of recombinant organisms. The development of molecular genetic tech-
niques has led to this relatively new field for addressing ecological
questions and issues.
monoclimax theory: The theory, promulgated by Frederic A. Clements,
that all communities within a given climatic region, despite initial dif-
ferences, eventually develop into the same climax community. See also
polyclimax theory.
monocropping: The common agricultural practice of planting and grow-
ing single crops, usually on a large tract of land, requiring the use of
chemical fertilizers and pest controls.
monoculture: A single crop used in monocropping.
monohybrid: An organism that is hybrid with respect to a single gene.
monsoon forest: Forest type found in tropical regions of the world where
there are annual periods of high rainfall.
mortality rate: The number of organisms in a population that die during a
given time interval.
Müllerian mimicry: Mimicry in which the mimic is toxic.
multiple use: Resource use in which land supports several concurrent
managed uses rather than single uses over time and space.
mutation: A change in the genetic sequence of an organism, sometimes
leading to an altered phenotype.
mutualism: A type of commensalism or symbiosis in which both sym-
biotes benefit from the association in terms of food, shelter, or protec-
tion.
mycobiont: The fungal part of a lichen.
712
Glossary
mycorrhiza (pl. mycorrhizae): A symbiotic relationship between a root and

a fungus in which the fungus lives either in or on the root and gains
food from it. The fungus increases absorption of water and minerals for
the root.
N: A standard abbreviation for the size of an actual population; if n̂, it is an

estimated value.
natality rate: Birthrate: the number of individuals that are born into a pop-
ulation during a given time interval.
natural selection: The process of differential survival and reproduction
that leads to heritable characteristics that are best suited for a particular
environment.
necroparasite: A parasite that consumes or lives in dead tissue.
nekton: An aquatic organism that has the ability to swim.
neoteny: Either the retention of immature characteristics in the adult form
or the sexual maturation of larval stages; it results in new kinds of adult
body plans.
neritic zone: The marine zone that begins at a depth of about 600 feet (180
meters), where the gradual slant of the continental shelf becomes a
sharp tilt toward the ocean floor, and extends to the ocean bottom.
nest parasite: Also called brood parasite; an individual (or species) that
lays its eggs in the nest of another individual (or species) and does no
parenting at all.
net primary productivity: The amount of energy, after plant respiration,
that is potentially available to primary consumers. See also gross pri-
mary productivity; secondary productivity.
neutral mutation: A mutation with no observable effect on the phenotype
of the cell or organism in which it occurs.
neutralism: A mutualism in which the two species are neither helped nor
harmed.
niche: An organism’s role in its habitat environment, such as food pro-
ducer, decomposer, parasite, plant eater (herbivore), meat-eater (carni-
vore). The sum of environmental conditions necessary for the survival
of a population of any species, including food, shelter, habitat, and all
other essential resources.
nitrogen cycle: The biogeochemical cycle of the element nitrogen.
nitrogen fixation: Process by which bacteria convert atmospheric nitrogen
to nitrogen-containing compounds, such as amino acids.
nocturnal: Active at night.
nomadic: Moving about from place to place according to the state of the
habitat and food supply.
713
Glossary
nonrandom mating: Mating that occurs whenever every individual does

not have an equal chance of mating with any other member of the popu-
lation.
nonrenewable resource: A natural resource, such as copper, coal, alumi-
num, and oil, that exists in a fixed amount and cannot be naturally re-
plenished at the rate it is being mined or removed.
nonruminating: Digesting grasses without chewing cud.
numerical response: The abundance of predators dependent upon the
abundance of prey in a habitat.
nutrient cycles, nutrient cycling: Large-scale movements of elements
(such as carbon, nitrogen, and water) through the living and nonliving
portions of an ecosystem.
old-growth forest: Ancient forests in which many trees are hundreds of

years old and which are among the richest ecosystems on earth. Con-
sidered nonrenewable.
omnivore: An animal that eats both plant material and animal material.
ontogeny: The successive stages during the development of an animal, pri-
marily embryonic but also postnatal.
opportunistic species: An invasive species that drives out native species
by competing with them for resources.
ordination: A method for collapsing community data for many species in
many communities along several environmental gradients onto a single
graph that summarizes their relationships and patterns.
organic: Living. At the molecular level, containing carbon atoms as a pri-
mary component.
organism: Any individual, self-contained life form, whether unicellular or
multicellular, whether protist, fungal, algal, plant, or animal.
orientation: An inherent sense of geographical location or place in time.
outbreeding: Interbreeding of stocks of a species that are unrelated to each
other. See also inbreeding.
outgroup: A group of organisms only distantly related to the groups being
examined in a cladistic study.
overfishing: Harvesting so many fish, including sexually immature fish,
that fishing becomes commercially inviable and ultimately leading to
the species’ extinction.
overgrazing: Destruction of vegetation by allowing too many grazing ani-
mals to graze beyond the carrying capacity of an area of rangeland.
overhunting: Harvesting so many terrestrial species (usually mammals or
reptiles), including sexually immature individuals, that the species is
ultimately threatened with extinction.
714
Glossary
ozone layer: The ozone-enriched layer of the stratosphere that filters out
some of the sun’s ultraviolet radiation, which causes skin and other
types of cancer.
P generation: Parental generation; the original individuals mated in a ge-

netic cross.
pair-bonding: Prolonged and repeated mutual courtship display by a mo-
nogamous pair, serving to cement the pair bond and to synchronize re-
productive hormones.
paleoecology: The study of past ecosystems and environments.
parasite: Any organism that lives on or in another living organism, the
host, and obtains its food from that host.
parasite-mix: All the individuals and species of symbiotes living concur-
rently in a host.
parasitism: The state and activities of being a parasite.
parasitoid: An insect, especially a wasp, that spends its larval stage in the
body of another insect and eats it, eventually emerging as a free-living
adult.
patch: A habitat fragment within a landscape, often occuring in a patch-
work of artificial and natural habitats strewn across a landscape in hap-
hazard and unplanned fashion.
pecking order: A dominance hierarchy in which the top individual can
threaten and force into submission any individual below it, the number
two individual can threaten anyone except number one, and the lowest-
ranked individual can threaten no one and must submit to everyone.
pelagic: The area of open water in the oceans; organisms that occur in the
water column.
per capita rate of increase: The difference between per capita births and
per capita deaths, defined as r. Also called the per capita rate of growth.
periodicity hypothesis: The proposal that mass extinctions have occurred
approximately every 26 million years over the past 250 million years.
permafrost: Permanently frozen layer of soil underlying tundra vegeta-
tion.
perturbation: Factors such as diseases, parasites, fire, and deforestation,
that disrupt ecosystems from within.
pesticide: Chemical or biological substances designed to kill unwanted
plants, fungi, or animals that interfere, directly or indirectly, with hu-
man activities. See also biopesticides.
phenology: The science that examines the relationships between climates
and climatic conditions and animal behaviors such as migrations or
parts of the life cycle such as flowering in plants.
715
Glossary
phenotype: The visible or outward expression of the genetic makeup of an

individual. See also genotype.
pheromone: A chemical produced by one member of a species that influ-
ences the behavior or physiology of another member of the same species.
phosphorus cycle: The biogeochemical movement of phosphorus through
an ecosystem.
photoperiodism: Regulation of a plant process, such as flowering, by the
relative length of light and dark periods in a twenty-four-hour day.
photorespiration: The process that occurs during photosynthesis when
the ratio of carbon dioxide to oxygen becomes too low. It results in the
production of less sugar than during regular photosynthesis, and some
carbon dioxide is formed.
photosynthesis: The process by which green plants and algae, called pri-
mary producers, use sunlight as energy to convert carbon dioxide and
water into energy-rich compounds such as glucose.
phyletic: related by or having a common ancestral line in evolutionary
terms.
phylogenetics: The study of the developmental history of groups of ani-
mals.
phylogeny: The evolutionary history of taxa, such as species or groups of
species; order of descent and the relationships among the groups are de-
picted.
physiological ecology: Sometimes called “autoecology,” “ecophysiology,”
or “comparative physiology,” a type of individual ecology that exam-
ines how life-forms function mechanically and physiologically in their
environments and how such factors as temperature, seasons, soil, and
nutrients affect survival and reproduction of those organisms. Analyzes
organismic adaptations from an engineering perspective in an evolu-
tionary context to determine the relationship between individuals’ per-
formance attributes, populations, and communities.
physiology: The study of the functions, activities, and processes of living
organisms.
phytophagous: Animals, also referred to as herbivorous, that feed on
plants.
phytoplankton: Small plants, often single-celled, that float in water. Phyto-
plankton in the ocean are responsible for much of the earth’s oxygen
production.
pioneer species: The earliest, hardy organisms that begin colonizing an
area in the first stage of ecological succession.
plankton: Small plants and animals that float freely in water; their small
size prevents them from having to swim to keep from sinking.
716
Glossary
plant ecology: The study of plant life (often including fungal and algal
forms) on all levels of ecology, from physiological to population to com-
munity to ecosystem ecology. Applications are particularly important
in agriculture.
poikilotherm: Cold-blooded or ectothermic; any organism having a body
temperature that varies with its surroundings; in general, reptiles, am-
phibians, fish, and invertebrates.
pollination: Transfer of pollen to a stigma in angiosperms or an ovule in
gymnosperms.
pollution ecology: The study of the impacts of water, air, and waste pollu-
tion on populations, communities, and ecosystems.
polyclimax theory: Within a given climatic region, there can be many cli-
maxes. See also monoclimax theory.
polygamy: A mating system in which one male mates with several females
(polygyny) or one female mates with several males (polyandry).
polygenic inheritance: Expression of a trait depending on the cumulative
effect of multiple genes; human traits such as skin color, obesity, and in-
telligence are thought to be examples of polygenic inheritance.
polymorphism: The occurrence of two or more structurally or behaviorally
different individuals within a species.
polyphyletic: Having similar characteristics but originating from more
than one ancestor.
polyploidy: Genetic condition in which the hereditary material is present
in three or more complete sets. Plants may be triploid (three sets of chro-
mosomes), tetraploid, or have even greater numbers.
population: A group of individuals of the same species that live in the
same location at the same time.
population analysis: The study of factors that influence growth of biologi-
cal populations.
population density: The number of individuals in a population per unit
area or volume.
population distribution: Variations in the density of a population in a par-
ticular area.
population dynamics: The patterns of population growth and decline over
the population’s existence, influenced by reproductive rates, predator-
prey relationships, carrying capacity, and other such factors.
population ecology: The study of the growth and decline of groups of in-
dividuals of the same species, and how these fluctuations function in re-
lation to other populations in the same ecosystem. Examines such fac-
tors as the availability of food and hence predation, herbivory, and
mutualisms. Community ecology is closely related to population ecol-
717
Glossary
ogy but focuses on the interactions between populations of different

species.
population fluctuations: Changes in population size over time.
population genetics: Branch of genetics that examines the movement of
genes through populations. A population geneticist might study ge-
netic drift, founder effect, or Hardy-Weinberg theorem.
population regulation: Stabilization of population size by factors such as
predation and competition, the relative impact of which depends on
abundance of the population in a habitat.
positive feedback loop: Situation in which a change in a certain direction
provides information that causes a system to change further in the same
direction. This can lead to a runaway or vicious cycle.
predation: The act of killing and consuming another organism. The organ-
ism that gains its nutrition in this way is the predator; the organism that
serves as the source of nutrition is the prey.
primary consumer: An organism that get its nourishment from eating pri-
mary producers, which are mostly green plants and algae.
primary metabolites: Sugar phosphates, amino acids, lipids, proteins, and
nucleic acids, all of which comprise the basic molecules necessary for a
cell to function.
primary producers: In an ecosystem, those organisms that form foods
from energy and simple chemical molecules.
primary succession: Succession in which the initial seral stage, or pioneer
community, begins on a substrate devoid of life or unaltered by living
organisms. See also secondary succession.
producers: See primary producers.
productivity: The rate of accumulation of biomass. See also gross primary
productivity; net primary productivity; secondary productivity.
promiscuity: A mating system in which sexual partners do not form last-
ing pair bonds; their relationship does not persist beyond the time
needed for copulation and its preliminaries.
protective mimicry: Use of both color and form to mimic an inanimate fea-
ture of the environment.
punctuated equilibrium: The idea that new species form during relatively
short speciation events (a few generations) and then persist for millions
of years in equilibrium (relatively unchanged) until they go extinct or
speciate again.
quadrat: A sample plot of a specific size and shape used in one method of
determining population size or species diversity.
718
Glossary
r strategy (r selection): A reproductive strategy involving high reproduc-

tive output; found often in unstable or previously unoccupied areas.
rain forest: Biome found in regions of the world where rainfall is high and
there are no dry periods. Rain forests occur in the tropics and along the
northwest coast of the United States.
random genetic drift: The random change of gene frequencies because of
chance, especially in small populations.
random mating: The assumption that any two individuals in a population
are equally likely to mate, independent of the genotype of either indi-
vidual; this is equivalent to saying that all the gametes of all the individ-
uals in a population are placed into a large pool, from which gametes
are paired at random.
rangeland: Open land of a wide variety of types, including grasslands, shrub-
lands, marshes, and meadows as well as some desert and alpine land.
recapture: See mark-capture-release methods.
reciprocal cross: A mating that is the reverse of another with respect to the
sex of the organisms that possess certain traits; for example, if a particu-
lar cross were tall male X short female, then the reciprocal cross would
be short male X tall female.
reciprocal relationship: Any type of coevolved, highly interdependent re-
lationship between two or more species.
reciprocal sacrifice: One explanation for acts of altruism among unrelated
animals; an individual sacrifice is made under the assumption that a
similar sacrifice may in turn aid the individual in the future.
recombinant DNA: DNA molecules that are the products of artificial re-
combination between DNA molecules from two different sources; im-
portant as a foundation of genetic engineering.
recombination: An exchange of genetic material, usually between two ho-
mologous chromosomes; provides one of the foundations for the ge-
netic reassortment observed during sexual reproduction.
red tide: An algal bloom that can be red, orange, brown, bright-green, or
even blooms that do not discolor the water in which they grow, usually
caused by dinoflagellates and often toxic.
reef: A carbonate structure that possess an internal framework that traps
sediment and provides resistance to wave action, thus forming habitat
for a rich and diverse marine community. Types of reefs include atoll,
barrier, fringing, and patch reefs.
reforestation: The growth of new trees in an area that has been cleared for
human activities, either occurring naturally or initiated by people.
relict population: A remnant population of otherwise extinct organisms.
replication: Copying of a DNA molecule, so that the two molecules formed
719
Glossary
are identical to each other and to the original molecule. The two mole-
cules formed are each composed of one strand from the original DNA
molecule and one newly synthesized strand.
reproductive (genetic) isolation: Describes any of many mechanisms that
prevent one organism from sexually reproducing with a second.
reproductive strategy: A set of traits that characterizes the successful re-
productive habits of a group of organisms.
reservoir host: A host species other than the one of primary interest in a
given research study.
resilience stability: Stability exhibited by a community that changes its
structure when disturbed but returns to its original structure when the
disturbance ends.
resistance: The ability of an organism, population, or community to sur-
vive in the face of natural or human-made threats.
resource: A requirement for life, such as space for living, food (for ani-
mals), or light (for plants), not including conditions such as tempera-
ture or salinity.
resource-holding potential: The ability of an individual to control a needed
resource relative to other members of the same species.
respiration: The utilization of oxygen; in air-breathing vertebrates, the in-
halation of oxygen and the exhalation of carbon dioxide.
restoration ecology: Restoration ecology is the study and implementation
of ways to return degraded or deteriorating communities and ecosys-
tems to their original condition. Restoration ecologists work to restore
habitat and return endangered species to viable numbers; they do not
seek to restore extinct species or re-create ancient habitats. See also con-
servation biology.
riparian ecosystem: An ecosystem in and around a river.
ritualization: An evolutionary process that formalizes the context and per-
formance of a display so that its meaning is clear and straightforward.
runoff: Surface water effluent from precipitation, irrigation, or other hu-
man activity and the chemicals and other materials it ultimately carries
into aquatic systems.
salinization: The accumulation of salts in soil.

saprophyte: Type of fungus or plant that gets its nutrition by digesting
dead plants. Many mushrooms grow as saprophytes.
saprovore: An organism that consumes dead or decaying plant or animal
matter.
savanna: Biome type characterized by widely spaced trees separated by
open, grassy regions.
720
Glossary
scale of being: An arrangement of life-forms in a single linear sequence

from lower to higher; also called a chain of being.
scavenger: An animal that feeds on the carcasses of other animals.
scrub community: Plant community characterized by stunted trees and
shrubs that may be widely spaced. Typical of poor soils.
secondary consumers: Carnivorous animals that eat herbivorous animals
(primary consumers).
secondary metabolite: A biochemical that is not involved in basic metabo-
lism, often of unique chemical structure and capable of serving a defen-
sive role for an organism.
secondary productivity: The rate at which animals produce their organic
matter by feeding on other organisms. See also gross primary productiv-
ity; net primary productivity.
secondary succession: Succession that starts in areas where an established
community has been disturbed or destroyed by natural forces or by hu-
man activities. See also primary succession.
seed dispersal: Process by which the seeds of a plant are distributed over a
wide area, away from the parent plant. Many seeds are adapted for dis-
persal by wind or animals.
selection: A process that prevents some individuals from surviving and
propagating while allowing others to do so. See also natural selection.
selective pressure: Evolutionary factors that favor or disfavor the genetic
inheritance of various characteristics of a species.
self-pollination: When pollen from the anther of a flower lands on the
stigma of the same flower.
semelparous species: Species, such as the Chinook salmon, that reproduce
only once before dying.
semiochemical: A chemical messenger that carries information between
individual organisms of the same species or of different species; phero-
mones and allelochemics are semiochemicals, but hormones are not.
sensitization: An arousal or an alerting reaction which increases the likeli-
hood that an organism will react; also, a synonym for loss of habituation
with increased intensity of response.
sequester: To store a material derived from elsewhere. In defenses, some
predators sequester defensive properties from their prey to defend
themselves from their own predators.
sex-role reversal: Generally used to refer to species in which the male does
most of the parenting.
sexual dimorphism: A difference in structure or behavior between males
and females.
sexual reproduction: Reproduction of cells or organisms involving the
721
Glossary
transfer and reassortment of genetic information, resulting in offspring

that can be phenotypically and genotypically distinct from either of the
parents.
sexual selection: The process that occurs when inherited physical or be-
havioral differences among individuals cause some individuals to ob-
tain more matings than others.
signal: Information transmitted through sound, such as bird calls, or
through sight, such as body posture.
signal pheromone: Nearly synonymous with releaser pheromone, but
used with mammals to remove the suggestion of a programmed re-
sponse and to indicate a more complex response.
sit-and-wait foraging: Sitting in one place, waiting, and attacking prey as
they move.
slash-and-burn (swidden) agriculture: An agricultural practice in which
forestland is cleared and burned for use in crop and livestock production.
SLOSS: An acronym for single large or several small preserves. This mir-
rors and reflects the current discussion as to how best preserve, protect,
and manage wildlife by use of one large park or preserve or several
smaller parcels.
social ecology: A branch of ecology, related to sociology, that critiques the
relationship between environmental destruction and social structure or
political ideology.
social grooming: An activity maintaining social interaction, whereby de-
bris is removed from a primate’s hair.
sociality: The tendency to form and maintain stable groups.
sociobiology: The study of the biological basis of the social behavior of an-
imals.
soil ecology: The study of soil as an ecosystem, including the interactions
of both abiotic and biotic components of soil: bacterial, fungal, plant
(humus, living roots), and animal (from protozoa and nematodes to in-
sects and earthworms). Soil ecology extends beyond the physical bor-
ders of soil to include the impact of soil on aboveground life-forms such
as larger plants and animals, as well as processes (geochemical cycles,
erosion, human agricultural practices) that impact soil. See also agricul-
tural ecology.
soil erosion: Loss of topsoil because of erosion due to runoff, winds, and
other forces.
soldiers: In insect societies, large workers that defend the colony and often
raid other colonies.
somatic hybrids: Plants that result from the fusion of two somatic (non-
reproductive) cells.
722
Glossary
specialists: Species with narrow niches that are not well generalized, often
able to live in only one habitat.
speciation: The evolution of new species as a result of geographic, phys-
iological, anatomical, or behavioral factors that prevent previously in-
terbreeding natural populations from breeding with each other any
longer.
species: A group of animals capable of interbreeding under normal natural
conditions; the smallest major taxonomic category.
species diversity: The variety of different organisms at the species taxo-
nomic level, a value that combines measures of both species richness
and species evenness.
species loss: Extinction.
species selection: The idea that species are independent entities with their
own properties, such as birth (speciation) and death (extinction); a
higher level of selection above that of natural selection is postulated to
take place on the species level.
species-specific: Innate and exclusive to a single species.
status badge: A visual feature that, based on its size or color or some other
variation, indicates the social status of the bearer.
stereotyped behavior: An unlearned and unchanging behavior pattern
that is unique to a species.
stimulus: Any environmental cue that is detected by a sensory receptor
and can potentially modify an animal’s behavior.
strategy: A behavioral action that exists because natural selection favored
it in the past (rather than because an individual has consciously decided
to do it).
stratification: Division of a community into layers, such as canopy, shrub,
and herb layers of a forest.
stress: A pressure on an organism, population, or community as a result of
ecosystem disturbance or an inadequate resource such as water or nu-
trients.
stromatolite: Fossilized masses of cyanobacteria that represent some of the
first evidence of photosynthetic life on earth.
subspecies: A group or groups of interbreeding organisms within a single
species that are distinct and separated from similar related groups but
not reproductively isolated.
substrate: The substance, such as soil or bark, on which a plant grows.
Also, in biochemical reactions, the chemical compound upon which an
enzyme acts.
succession: Change in a plant or animal community over time, with one
kind of organism or plant being replaced by other organisms or plants
723
Glossary
in a more or less predictable pattern. See also primary succession; sec-

ondary succession.
superorganism concept: An “organism” composed of more than one mem-
ber, such as an insect society, reflecting the remarkable degree of coordi-
nation between individuals.
survivorship: The pattern of survival exhibited by a cohort throughout its
life span.
sustainable development: The study and implementation of methods of
agriculture, mining, timber harvesting, and other human interactions
with the natural environment in a way that ensures minimal or no impact
on the original ecosystems, to the end of conserving natural resources
and minimizing impact on the organisms that occupy the habitats in-
volved. The term “sustainable” is often coupled with more specific dis-
ciplines: “sustainable agriculture,” “sustainable forestry,” and so on. See
also restoration ecology.
swidden agriculture: See slash-and-burn agriculture.
switching: The phenomenon that occurs when a predator has a choice of
several prey species and learns to prefer one of them over a previous
prey; if the preferred prey is sufficiently abundant, the predator will be-
gin to concentrate on the preferred prey and may change its searching
and other prey-related behaviors.
symbiont: A member of a symbiotic relationship.
symbiosis: A type of coevolved relationship between two species in which
both participants benefit; a type of mutualism.
sympatric: Living in the same place, not separated by a barrier that would
prevent interbreeding.
sympatric speciation: Evolution of two groups into separate species while
living in overlapping geographic locations.
synergism: The result of the interactions of a number of agents, operations,
organisms, and other factors such that the final effect is greater than the
sum of the individual effects.
systematics: The subdivision of biology that deals with the identification,
naming, and classification of organisms and with understanding the
evolutionary relationships among them. See also classification; taxon-
omy.
taiga: Biome located across Canada, northern Europe, and northern Asia
that consists of forests of spruces, firs, and birches. See also boreal forest.
taxonomy: A classification scheme for organisms based primarily on struc-
tural similarities; taxonomic groups consist of genetically related ani-
mals. See also classification; systematics.
724
Glossary
temperate deciduous forest: Biome located in eastern North America

south of the taiga, central Europe, and eastern China, characterized by
forests of tree species, most of which lose their leaves every autumn.
temperate mixed forest: A subdivision of the temperate deciduous forest
in which pines share importance with deciduous trees. These forests are
usually found on sandy soils in the southeastern United States.
temporal variation: Variation across time.
terrestrial: Living on land.
territorial behavior: The combination of methods and actions through
which an animal or group of animals protects its territory from invasion
by other species.
threat display: A territorial behavior exhibited by animals during defense
of a territory, such as charging, showing bright colors, and exaggerating
body size.
threatened species: Animals or plants, as designated by official bodies
such as the U.S. Fish and Wildlife Service, whose members are so few in
number that they may soon become endangered and then extinct.
topography: The structure and configuration of the earth’s surface, includ-
ing its natural and human-made features.
toxin: Any substance, such as the venom in snakes or spiders, that is toxic
to an animal.
trace element: Chemical element required in very small quantities for nu-
trition.
transgenic: Possessing one or more genes from another organism, a con-
cept important for the study of genetic mutations.
transpiration: Evaporation of water from the leaves and stems of plants.
Most transpiration occurs through open stomata.
trophic levels: Positions at which specific organisms obtain their nutrition
within a food chain. Plants and photosynthetic algae, which are pri-
mary producers, occupy the first trophic level, followed by primary and
secondary consumers, scavengers, and decomposers.
tropical rain forest: Biome found in tropical regions throughout the world
that experience rainfall year-round. These are the most biodiverse eco-
systems, often containing thousands of species of plants in a single acre.
tropism: Plant growth response to an external stimulus. The plant may
grow toward or away from the stimulus. Includes geotropism, gravi-
tropism, heliotropism, phototropism, and thigmotropism.
tundra: Regions where no trees grow because of frozen soil or extreme
water runoff due to steep grades (at high altitudes) are known as tun-
dra. See also alpine tundra; Arctic tundra.
725
Glossary
understory: In a rain forest, the shorter trees and shrubs that grow under
and live in the shade of the canopy formed by the crowns of the tallest
trees.
uniformitarianism: The belief that the earth and its features are the result
of gradual biological and geological processes similar to the processes
that exist today.
urban ecology: The study of the ecology of cities and human settlements,
including energy and water flows, resource use, and how and where
humans build. See also landscape ecology.
urban heat island: A relatively hot area in the atmosphere above an urban
area produced by a concentration of cars, factories, reflective surfaces,
and other heat-producing factors.
urbanization: The process of converting natural habitats into urban com-
plexes.
vector: An organism, such as a baterium, a mosquito, a virus, or a rodent or

other mammal, that transmits pathogens from one host to another.
veld: A grassland ecosystem, especially in southern Africa.
venom: A toxic substance that must be injected (instead of ingested) to im-
mobilize or kill prey.
virion: A virus particle consisting of genetic material surrounded by a pro-
tein coat.
viroid: Small, viruslike infectious molecules of RNA without protein coats.
virus: A microscopic infectious particle composed primarily of protein and
nucleic acid; bacterial viruses, or bacteriophages, have been important
tools of study in the history of molecular genetics.
visual predation: Catching prey (such as insects) by sighting them visu-
ally, judging their exact position and distance, and pouncing on them.
warm-blooded: Referring to animals whose body temperatures are main-

tained at a constant level by their own metabolisms.
warning coloration: The bright colors seen on many dangerous and unpal-
atable organisms that warn predators to stay away.
weeds: hardy plants, usually invasive species, that successfully compete
with native plants to the latter’s eventual weakening and extinction.
wetlands: Transitional areas between aquatic and terrestrial habitats, home
to a variety of flood-tolerant and salt-tolerant species.
wildlife: All living organisms; traditionally, the term included only mam-
mals and birds that were hunted or considered economically important.
workers: Sterile, wingless female social insects.
726
Glossary
zonation: The distribution of organisms and other elements of ecosystems

into regular or distinct biogeographical zones.
zoogeography: The study of the distribution of animals over the earth.
zoology: The study of the classification, anatomy, and physiology of ani-
mal life.
zooplankton: Small animals, often single-celled, that float or swim weakly.
zygote: A diploid cell produced by the union of a male gamete (sperm)
with a female gamete (egg); through successive cell divisions, the zy-
gote will eventually give rise to the adult form of the organism.
727
WEB SITES
NOTE: The Web sites listed below offer an entry into the many resources available
on the Internet. These are simply some of the better known and more reliable sites.
The editors accessed these URLs in April, 2003.
Biodiversity and Ecosystem Function Online

http://www.abdn.ac.uk/ecosystem/bioecofunc
An “informal collaboration between several research institutes within the
United Kingdom that share similar research interests. Hosted by the
University of Aberdeen, Scotland, the site aims to be a resource for
researchers specifically concerned with how the richness of species
(or functional groups) affects ecosystem function. A list of the partici-
pants and a link to their respective research interests can be accessed
through the ‘Project Partners’ page.” Offers a bibliography on ecosys-
tem ecology.
British Ecological Society

http://www.britishecologicalsociety.org
The BES “is an active and thriving organisation with something to offer
anyone with an interest in ecology. Academic journals, teaching re-
sources, meetings for scientists and policy makers, career advice and
grants for ecologists” are among its activities. The Web site offers pages
for general interest, students, teachers (including access to curriculum-
driven publications), ecological issues, and career information.
Earth’s 911
http://www.earth911.org/master.asp
Aimed at children, students, and the general public, this site offers in-
formation on recycling and waste management options, including links
to state-by-state listings of recycling centers, businesses, and regula-
tions.
Ecological Society of America

http://www.esa.org
A nonpartisan, nonprofit organization of scientists founded in 1915 to
“promote ecological science by improving communication among ecol-
ogists; raise the public’s level of awareness of the importance of ecologi-
cal science; increase the resources available for the conduct of ecological
science; and ensure the appropriate use of ecological science in environ-
729
Web Sites
mental decision making by enhancing communication between the eco-

logical community and policy-makers.” Sponsors the Sustainable Bio-
sphere Initiative. Offers an Education Section as well as resources for
career placement, publications, and links to other ecology sites.
Ecology WWW Page

http://www.botany.net/Ecology
Maintained by Anthony R. Brach of the Missouri Botanical Garden and
Harvard University Herbaria, this list of links is maintained as a volun-
teer service to students, teachers, researchers, and others interested in
the science of ecology.
Envirolink
http://www.envirolink.org
This nonprofit organization has been providing access to online environ-
mental resources since 1991, organized by categories such as Ecosys-
tems, Environmental Disasters, Forests, and many more. The Ecology
subsection files sites under such subcategories as Actions You Can Take,
Educational Resources, General Info, Government Resources, Organi-
zations, Jobs and Volunteer Opportunities, and Publications.
Environmental Protection Agency

http://www.epa.gov
This U.S. government agency offers information and Web links organized
by ecosystems such as the following: aquatic ecosystems, coasts, coral
reefs, deserts, ecological assessment, ecological monitoring, ecological
restoration, endangered species, environmental indicators, estuaries,
exotic species, forests, freshwater ecosystems, lakes, landscape ecology,
marine ecosystems, oceans, species, terrestrial ecosystems, urban eco-
systems, watersheds, and wetlands.
National Academy of Sciences

http://www4.nationalacademies.org/nas/nashome.nsf
A “private, nonprofit, self-perpetuating society of distinguished scholars
engaged in scientific and engineering research, dedicated to the further-
ance of science and technology and to their use for the general welfare.
Upon the authority of the charter granted to it by the Congress in 1863,
the Academy has a mandate that requires it to advise the federal gov-
ernment on scientific and technical matters.”
730
Web Sites
National Council for Science and the Environment

http://www.ncseonline.org
A U.S. government-sponsored Web site which links to the National Library
for the Environment. From here, students can search more than 1,200
congressional reports on topics in agriculture, biodiversity, climate
change, energy, forests, pesticides, pollution, public lands, stratospheric
ozone, wetlands, and a host of other environmental topics.
National Oceanic and Atmospheric Administration

http://www.noaa.gov
NOAA maintains pages on oceans, satellites, fisheries, weather, climate,
coasts, and current events broken down by type of environmental im-
pact: dust, fires, floods, ice, oil spills, snow, severe weather, tropical, and
volcanic. In addition, NOAA’s library, photo library, other pages on top-
ics of ecological interest (such as coral reefs and El Niño), and organized
lists of links provide access to other sites and research.
National Resources Conservation Service

http://www.nrcs.usda.gov
The mission of this division of the U.S. Department of Agriculture is to “as-
sist owners of America’s private land with conserving their soil, water,
and other natural resources.” The NRCS maintains two pages of inter-
est: “Ecological Sciences” for agribusiness professionals and a page for
teachers and kids with educational materials on soils, conservation bi-
ology, composting, and the like.
The Need to Know Library: Ecology and Environment Page

http://www.peak.org/~mageet/tkm/ecolenv.htm
Lists topics “under three headings: Ecology, Conservation Biology, and
Botany and Systematics. Links to home pages of botanical and ecolog-
ical societies are provided in the Journals and Societies List. The Eco-
logical Software List offers links to sites describing software applica-
tions for analyzing or organizing ecological data. The Natural Resource
Agencies List provides links to home pages of government agencies
that conduct ecological research or that are responsible for the manage-
ment of various ecosystems. The Natural Areas and Reserves heading
lists links to sites that feature information about natural areas, LTER
sites, national parks, and wilderness areas. The Environment List refers
to sites dealing with ecosytem conditions and the Environmental Soci-
eties List is a catalog of links to home pages of environmental organiza-
tions.”
731
Web Sites
Sierra Club
http://www.sierraclub.org
In addition to its outreach programs encouraging public involvement in
recycling, conservation, and nature activities, the Sierra Club is active in
environmental issues, offering updates on current events in water, en-
ergy, global population, and more.
Society for Conservation Biology

http://conbio.net
The Virtual Library of Ecology and Biodiversity is maintained through this
site.
UNEP-WCMC Protected Areas Information

http://www.wcmc.org.uk/data/database/un_combo.html
Sponsored by the United Nations Environment Programme and the World
Conservation, this page provides access to information on nearly all na-
tions of the world, including basic facts, history, and present policies
concerning protected areas. Each essay is lengthy and detailed and in-
cludes references.
United National Environment Programme

http://www.unep.org
UNEP’s mission is “to provide leadership and encourage partnership in
caring for the environment by inspiring, informing, and enabling na-
tions and peoples to improve their quality of life without compromising
that of future generations.” Its Web site offers resources and links on
habitats, species, regions, climate change, protected areas, and much
more—including a kids’ page that offers facts and educational games
on the atmosphere, forests, biodiversity, polar regions, whales, and
aquatic ecosystems.
U.S. Geological Survey

http://www.usgs.gov
Not simply a geology resource, the USGS maintains a wealth of data and
fact sheets on biological and ecological information as well, searchable
by topic or state. Topics include Biological Resources, Earthquakes,
Floods, Ground Water, Maps, Streamflow, Water Quality, and more.
732
Web Sites
USDA Forest Service

http://www.fs.fed.us
This U.S. government site offers information on public lands in national
forests and grasslands, organized state by state, as well as a photo gal-
lery, maps, and information on the programs overseen by the agency.
World Conservation Union

http://www.iucn.org
Known also by its former initials IUCN, the World Conservation Union
“seeks to influence, encourage and assist societies throughout the
world to conserve the integrity and diversity of nature and to ensure
that any use of natural resources is equitable and ecologically sustain-
able.” Provides access to library catalog, publications, programs, con-
tacts.
World Resources Institute

http://www.wri.org
A nonprofit environmental research and policy organization that works
with governments, the private sector, and civil society groups in more
than one hundred countries around the world. Offers updates on glo-
bal environmental and ecological issues. Maintains a section called
“EarthTrends” in which maps, data tables, searchable databases, and
country profiles can be accessed by topic: Coastal and Marine Eco-
systems; Water Resources and Freshwater Ecosystems; Climate and
Atmosphere; Population; Health and Human Well-being; Economics;
Business and the Environment; Energy and Resources; Biodiversity
and Protected Areas; Agriculture and Food; Forests, Grasslands and
Drylands.
733
CATEGORIZED INDEX
Agricultural ecology Reproductive strategies, 576
Biopesticides, 65 Territoriality and aggression, 633
Erosion and erosion control, 211
Genetically modified foods, 284 Biomes
Grazing and overgrazing, 304 Biomes: determinants, 55
Integrated pest management, 351 Biomes: types, 59
Multiple-use approach, 422 Chaparral, 76
Pesticides, 470 Deserts, 154
Rangeland, 560 Forests, 269
Slash-and-burn agriculture, 590 Grasslands and prairies, 298
Soil, 594 Habitats and biomes, 313
Lakes and limnology, 364
Aquatic and marine ecology Marine biomes, 391
Acid deposition, 1 Mediterranean scrub, 399
Lakes and limnology, 364 Mountains, 419
Marine biomes, 391 Old-growth forests, 452
Ocean pollution and oil spills, 444 Rain forests, 549
Reefs, 564 Rain forests and the atmosphere,
554
Behavioral ecology Rangeland, 560
Altruism, 18 Reefs, 564
Communication, 95 Savannas and deciduous tropical
Competition, 111 forests, 586
Defense mechanisms, 125 Taiga, 629
Displays, 167 Tundra and high-altitude biomes,
Ethology, 215 655
Habituation and sensitization, 319 Wetlands, 672
Herbivores, 326
Hierarchies, 329 Chemical ecology
Insect societies, 343 Allelopathy, 15
Isolating mechanisms, 358 Bioluminescence, 43
Mammalian social systems, 385 Communication, 95
Migration, 407 Defense mechanisms, 125
Mimicry, 415 Genetically modified foods, 284
Omnivores, 455 Metabolites, 402
Pheromones, 476 Pheromones, 476
Poisonous animals, 486 Poisonous animals, 486
Predation, 536 Poisonous plants, 490
735
Categorized Index
Community ecology Biomes: determinants, 55

Allelopathy, 15 Biomes: types, 59
Animal-plant interactions, 24 Chaparral, 76
Biodiversity, 32 Communities: ecosystem
Biogeography, 37 interactions, 100
Biological invasions, 40 Desertification, 149
Coevolution, 86 Deserts, 154
Communities: ecosystem Ecosystems: definition and
interactions, 100 history, 184
Communities: structure, 104 Ecosystems: studies, 191
Competition, 111 Erosion and erosion control, 211
Food chains and webs, 255 Forest fires, 258
Gene flow, 274 Forests, 269
Genetic diversity, 278 Genetic diversity, 278
Lichens, 381 Geochemical cycles, 288
Mycorrhizae, 425 Grasslands and prairies, 298
Pollination, 495 Habitats and biomes, 313
Speciation, 604 Hydrologic cycle, 338
Succession, 612 Invasive plants, 354
Symbiosis, 621 Isolating mechanisms, 358
Trophic levels and ecological Lakes and limnology, 364
niches, 641 Marine biomes, 391
Mediterranean scrub, 399
Ecoenergetics Mountains, 419
Balance of nature, 28 Nutrient cycles, 440
Biomass related to energy, 50 Old-growth forests, 452
Food chains and webs, 255 Rain forests, 549
Geochemical cycles, 288 Rain forests and the atmosphere,
Herbivores, 326 554
Hydrologic cycle, 338 Rangeland, 560
Nutrient cycles, 440 Reefs, 564
Omnivores, 455 Savannas and deciduous tropical
Phytoplankton, 482 forests, 586
Rain forests and the atmosphere, Soil, 594
554 Taiga, 629
Trophic levels and ecological Trophic levels and ecological
niches, 641 niches, 641
Tundra and high-altitude biomes,
Ecosystem ecology 655
Biodiversity, 32 Wetlands, 672
Biological invasions, 40
736
Categorized Index
Ecotoxicology Isolating mechanisms, 358

Acid deposition, 1 Natural selection, 428
Biological invasions, 40 Nonrandom mating, genetic drift,
Biomagnification, 47 and mutation, 435
Biopesticides, 65 Paleoecology, 464
Deforestation, 131 Punctuated equilibrium vs.
Eutrophication, 222 gradualism, 543
Genetically modified foods, 284 Speciation, 604
Integrated pest management, 351 Species loss, 608
Invasive plants, 354
Ocean pollution and oil spills, Global ecology
444 Biodiversity, 32
Ozone depletion and ozone holes, Biomes: determinants, 55
457 Biomes: types, 59
Pesticides, 470 Biosphere concept, 69
Phytoplankton, 482 Geochemical cycles, 288
Pollution effects, 500 Global warming, 292
Slash-and-burn agriculture, 590 Greenhouse effect, 308
Soil contamination, 601 Hydrologic cycle, 338
Waste management, 667 Ozone depletion and ozone holes,
457
Evolutionary ecology Rain forests and the atmosphere,
Adaptations and their 554
mechanisms, 7
Adaptive radiation, 12 History of ecology
Coevolution, 86 Ecology: history, 179
Colonization of the land, 90 Ecosystems: definition and
Convergence and divergence, 120 history, 184
Dendrochronology, 145 Evolution: history, 236
Development and ecological
strategies, 161 Landscape ecology
Evolution: definition and theories, Landscape ecology, 374
227 Urban and suburban wildlife, 659
Evolution: history, 236
Evolution of plants and climates, Paleoecology
241 Colonization of the land, 90
Extinctions and evolutionary Dendrochronology, 145
explosions, 246 Evolution: definition and theories,
Gene flow, 274 227
Genetic drift, 281 Evolution of plants and climates,
Genetically modified foods, 284 241
737
Categorized Index
Extinctions and evolutionary Restoration and conservation

explosions, 246 ecology
Paleoecology, 464 Biodiversity, 32
Conservation biology, 119
Physiological ecology Deforestation, 131
Adaptations and their Endangered animal species, 196
mechanisms, 7 Endangered plant species, 205
Bioluminescence, 43 Erosion and erosion control, 211
Camouflage, 72 Forest management, 263
Defense mechanisms, 125 Genetic diversity, 278
Mimicry, 415 Grazing and overgrazing, 304
Pheromones, 476 Integrated pest management, 351
Poisonous animals, 486 Multiple-use approach, 422
Poisonous plants, 490 Old-growth forests, 452
Pollination, 495 Reforestation, 572
Tropisms, 650 Restoration ecology, 583
Soil contamination, 601
Population ecology Species loss, 608
Adaptive radiation, 12 Sustainable development, 618
Biodiversity, 32 Urban and suburban wildlife, 659
Biogeography, 37 Waste management, 667
Clines, hybrid zones, and Wildlife management, 677
introgression, 80 Zoos, 681
Convergence and divergence, 120
Demographics, 137 Soil ecology
Extinctions and evolutionary Erosion and erosion control, 211
explosions, 246 Soil, 594
Gene flow, 274 Soil contamination, 601
Genetic diversity, 278
Genetic drift, 281 Speciation
Human population growth, 333 Adaptive radiation, 12
Insect societies, 343 Clines, hybrid zones, and
Nonrandom mating, genetic drift, introgression, 80
and mutation, 435 Convergence and divergence, 120
Population analysis, 507 Evolution: definition and theories,
Population fluctuations, 513 227
Population genetics, 520 Evolution of plants and climates,
Population growth, 528 241
Punctuated equilibrium vs. Extinctions and evolutionary
gradualism, 543 explosions, 246
Reproductive strategies, 576 Gene flow, 274
738
Categorized Index
Isolating mechanisms, 358 Biomes: determinants, 55

Natural selection, 428 Biomes: types, 59
Punctuated equilibrium vs. Biosphere concept, 69
gradualism, 543 Deep ecology, 123
Speciation, 604 Ecology: definition, 171
Ecosystems: definition and
Theoretical ecology history, 184
Balance of nature, 28 Sustainable development, 618
739
SUBJECT INDEX
Abiotic components of 287; and rangelands, 562; slash-
ecosystems, 186, 191, 314 and-burn, 590, 593; world food
Abundance, cycles of, 513, 539 supplies, 336
Abyssal marine zone, 63, 392 Ailanthus (tree-of-heaven), 659
Acacia-ant mutualisms, 87 Air pollution, 591; lichens as
Acceleration (heterochrony), 163 bioindicators, 383
Acid deposition, 1, 6, 110, 189, 591, Alarm calls, 97
631 Alarm pheromones, 476
Acid rain, 501; and soil Alerting system, 321
generation, 594 Algae, 315; earliest life on land, 91;
Adaptation, 171, 428, 521; eutrophic environments, 223;
behavior as, 217; Darwin on, killer, 357; phytoplankton, 482.
238; mechanisms of, 7, 11; See also Phytoplankton;
migration, 408; ontogeny as, Plankton
162; and speciation, 606; Algal blooms, 224, 370, 482-483
Triassic period, 250 Aliphatics (hydrocarbons), 444
Adaptive coloration, 10 Alkaloids, 103, 403, 491
Adaptive radiation, 12, 14, 122, Alleles, 12, 274, 281, 435;
239; and speciation, 606 frequency, 436
Adders, 488 Allelochemicals, 15
Adelgids, 41 Allelopathy, 15, 17, 101, 354
Adult survivorship patterns, 139 Allen’s rule, 81
Africa, habitat loss, 207 Alligator River Wildlife Refuge,
Agave lechuguilla (indicator 116
species), 156 Alligators, 203
Age structures, 140, 508; Allochthonous deposition, 465
population analysis, 511 Allogenic clastic materials, 366
Aggregation pheromones, 476 Allomones, 477
Aggression, 22, 633, 640; in Allopatric speciation, 543, 604
dominance hierarchies, 329 Alpha individuals, 330
Aggressive mimicry, 417, 538 Alpine tundra, 61, 315, 655
Agrichemicals, 500 Altruism, 18, 23, 522
Agricultural ecology. See Alvarez, Luis, 247
Agriculture; Categorized Index Alvarez, Walter, 247
Agriculture; uses of allelopathy, Amazon basin, 558
15; and deforestation, 131; Amber, 466
genetic diversity, 279; Ambush predation, 539
genetically modified crops, 284, Amensalisms, 100
741
Subject Index
American Acclimitization Society, Arabian Gulf, 450

662 Araceae (philodendron), 491
American pronghorns, 300 Arbor Day, 575
American Society of Limnology Arbuscules, 427
and Oceanography, 180 Arbutoid mycorrhizae, 426
Anaerobic bacteria, 369 Archaeocalamites, 242
Anaphylactic shock, 487 Archaeocyathids, 567
Anaximander, 236 Arctic terns, migration, 411
Ancient Bristlecone Pine Forest, Arctic tundra, 61, 314, 655
146 Argyroxiphium sandwicense
Anemones, 624 (silversword), 13
Angiosperm evolution, 86, 244 Arid climates, 149, 153
Anglerfish, 396 Aromatics (hydrocarbons), 445
Animal-plant interactions, 10, 24, Arrestant pheromones, 476
27, 100, 105; herbivores, 326; Artemisia tridentata (indicator
pollination, 86 species), 156
Anoxia, eutrophic environments, Arthropods, 487
224 Artiodactyls, 326
Antennae and pheromones, 478 Asexual reproduction, 524, 605
Anthocyanidins, 405 Aspens, quaking, 629
Anthocyanins, 405 Aspirin, 405
Antibiotic resistance, 526 Associational defenses, 128
Antibiotics, 15 Aster family, 13, 157
Antithesis principle, 168 Atlantic Ocean, surface
Ants, 343; and aphids, 128; temperatures, 150
competition among, 115; Atmosphere, 69; ancient, 556;
mutualisms with acacias, 87, impact of desertification, 151;
174; pheromones, 477 Devonian oxygenation, 93;
Aphids, 343; and ants, 128 ozone, 556; ozone depletion,
Aphotic marine zone, 391 457, 463; rain forests’ impact,
Aplysia (sea hare), 321 554, 559
Aposematic coloration, 128, 167 Atoll reefs, 564
Aposematism, 538 Atomic Energy Commission, 180,
Aquatic and marine ecology. See 188
Categorized Index Atropine, 404
Aquatic biomes, 315 Attractants; phenolics, 405;
Aquatic communities, 173 pheromones, 476
Aquatic ecosystems, 614; nutrient Auditory calls; communication, 97;
sources, 440; production in, 188 displays, 169
Aquatic plants; invasive, 357; Audubon, John James, 198
wetlands, 674 Audubon Society, 678
742
Subject Index
Australia, habitat loss, 207 Bats; desert, 157; as pollinators,

Australian honey eaters, 636 497
Autochthonous deposition, 465 Beaded lizards, 486
Autotomy (defense mechanism), Beagle, voyage of the, 238
129 Bears, 13
Autotrophs, 50, 86, 111. See also Beebe, William, 197
Producers Beefalo, 84
Auxins, 650 Beer, Gavin de, 163
Bees, 344; Africanized, 116; dance
Baboons; competition among, 114; language, 215; as pollinators,
hierarchies, 330; social systems, 496
386 Beetles; bark, 511; bombardier,
Bacillus cereus (biopesticide), 66 126; as pollinators, 497
Bacillus popilliae (biopesticide), 66 Behavior; as camouflage, 74; and
Bacillus thuringiensis communication, 95; ethology,
(biopesticide), 66, 285 215; and reproductive
Backcrossing, 83 strategies, 576
Background extinctions, 246 Behavioral adaptations, 10
Bacteria; anaerobic, 224; Behavioral displays, 167
bioluminescent, 43; Behavioral ecology, 218. See also
biopesticides, 65; Categorized Index
cyanobacteria, 225, 393, 465, Behavioral genetics, 217
482; as decomposers, 112, 185, Behavioral isolation, 358
290, 368; in digestion, 102, 305; Behaviorism, 216
mutation rates in, 438; nitrogen Benthic marine zone, 391, 397
fixation, 289; oil-eating, 448; Bergmann’s rule, 80
phosphorus cycle, 290; in soil, Beta-carotene, 405
316, 596; symbiotic, 127; Beta individuals, 330
weathering agents, 173 Bighorn sheep, 157
Bacterioplankton, 392-393 Bioaccumulation. See
Badgers, defense mechanisms of, Biomagnification
126 Biodiversity, 32, 36, 134, 618; and
Baer, Karl Ernst von, 161 adaptation, 7; ancient, 251; and
Baer’s laws, 161 biogeography, 37; and
Balance of nature, 28, 31, 255, 517 pollution, 500; rain forests, 558;
Bald eagles, 199 urban and suburban wildlife,
Barnacles, 623 660
Barrier reefs, 564 Biogenetic law, 162
Base flow, 341 Biogeochemical cycles; hydrologic
Batesian mimicry, 128, 416, 538 cycle, 338, 342; nutrient cycles,
Bathypelagic marine zone, 392 440, 443. See also Carbon cycle;
743
Subject Index
Geochemical cycles; 497; population fluctuations,

Hydrologic cycle; Nitrogen 515; territorial behavior, 633;
Cycle; Nutrient cycles; Oxygen urban habitats, 660
cycle; Phosphorus cycle Birthrates (population analysis),
Biogeography, 37, 39, 179, 233; 137, 530
islands, 33 Bison, 300, 609; introgression of,
Bioherms, 564 84; overhunting of, 198
Biological diversity, 612 Black-collared lizards, 158
Biological invasions, 40, 42 Black-footed ferrets, 109, 203, 300
Biological phenomena diversity, Black mambas, 488
35 Black widow spiders, 158, 487
Bioluminescence, 43, 46, 96, 392 Blackbirds, competition among,
Biomagnification, 47, 49, 474 115
Biomass, 50, 54, 174, 186-187, 256, Blight, chestnut, 355
612; grasslands, 298 Blood flukes, 625
Biomes, 313, 318; determinants of, Bobcats, 157
55, 58; and habitats, 314; Body size; and clines, 80; and
studies, 181; table, 56; types, 59, reproduction, 576
64. See also Alpine tundra; Bogs, 673
Arctic tundra; Boreal forests; Bombardier beetles, 126
Chapparal; Deciduous tropical Boreal forests, 56, 60, 270, 629, 632,
forests; Deserts; Ecosystems; 656
Ecotones; Forests; Grasslands; Bormann, F. Herbert, 193
Lake ecosystems; Marine Bottleneck effect on population
biomes; Mediterranean scrub; size, 437
Mountain ecosystems; Old- Boyce, Mark, 579
growth forests; Prairies; Rain Brachiopods, 248
forests Brachystegia (Central Africa), 586
Biopesticides, 65, 68, 355, 470; in Brackish marshes, 674
integrated pest management, Bradley, Richard, 28
352 Branching evolution, 228, 237
Biosphere, 69, 71, 191, 316 Breeding cycles, 514
Biostromes, 564 Breeding programs, 480
Biotic components of ecosystems, Bristlecone pines, 146
186, 192 British Ecological Society, 180
Biotic potential, 530 Brown pelicans, 203
Birches, 629 Brown tides, 224. See also Algal
Birdcalls and birdsongs, 97; and blooms; Eutrophication;
territoriality, 634 Phytoplankton; Red tides
Birds; communication, 96; Bryozoans, 567
omnivorous, 456; pollinators, Buffalo, 609; overhunting of, 198
744
Subject Index
Buffon, Comte de, 29 326; marine, 395; social

Bullfrogs, 158 organization, 389
Bureau of Land Management, 677 Carnivorous plants, 208
Burgess Shale, 249 Carr, Archie, 408
Buri, Peter, 438 Carrying capacity, 81, 187, 301,
Buteo jamaicensis (red-tailed 304, 410, 511, 531, 577, 678
hawk), 663 Carson, Rachel, 30, 193, 200
Butterflies, 203; monarch, 538; as Castes and social insects, 345
pollinators, 497; viceroy, 538 Castor bean plant, 492
Cat hierarchies, 331
Caatinga (Amazon basin), 586 Catastrophism, 238
Cacti, 156; convergent evolution, Cations, 595
120; endangered, 208 Cedar Bog Lake, Minnesota, 186,
Caffeine, 404 191
Calcareous algae, 567 Cellular respiration, 554
California, fire management in, Cellulose, 102
400 Cenozoic era, 244
California condors, 203 Censuses (demographic tool), 142
Calvin cycle, 556 Central American dry forests, 588
Cambrian explosion, 249 Central Park, New York City, 663
Camouflage, 72, 75, 167; and Ceratius, 396
bioluminescence, 45; vs. Cerrado (Brazil), 586
mimicry, 415; peppered moth, CFCs. See Chlorofluorocarbons
10 Challenger, HMS, 182
Canada, acid precipitation in, 631 Chambers, Robert, 237
Canada thistle, 356 Channel Islands, California, 206
Cancer medications, 404 Chaparral, 62, 76, 79, 399
Canopy (rain forests), 269, 551, Character displacement, 644
558 Charophytes, 91
Captive breeding, 684 Chemical ecology. See
Carbamates, 471 Categorized Index
Carbon cycle, 288, 441, 483; rain Chen hyperborea (snow goose), 435
forests, 556 Chenopodiaceae (indicator species),
Carbon dioxide; and global 157
warming, 310; rain forests, 554 Chestnut trees, 356
Carboniferous period, 242 Chihuahuan Desert, 156
Carew, Thomas J., 322 Chimpanzees, habituation studies,
Carnegiea gigantecus (indicator 324
species), 156 Chlorinated hydrocarbons, 470
Carnivores, 111, 174, 256, 455, 561, Chlorofluorocarbons, 308, 459, 501
641; bears, 13; and herbivores, Circadian rhythms, 409
745
Subject Index
Circumnutation, 652 Columella cells, 651

Clean Water Act (1965), 675 Commensalisms, 87, 100, 622
Cleaning symbiosis, 624 Communication, 95, 99; bees, 215,
Clear-cutting, 53, 266, 334, 552; 345; and bioluminescence, 44;
reforestation, 572 insects, 343, 345; and
Clements, Frederic E., 30, 182, pheromones, 480
615 Communities; ecosystem
Cleveland, Grover, 422 interactions, 100, 103; and
Climate change, global warming, ecosystems, 184; and habitats,
294 314; isolated, 37; lentic and
Climates; and the biosphere, 70; lotic, 315; structure, 104, 110,
and tree rings, 145 612; succession, 612, 617
Climax communities, 612, 617 Community ecology, defined, 173.
Climax pattern concept, 615 See also Categorized Index
Climax stage, 173 Compartmentalization, 104
Clines, 80, 85 Compass plants, 653
Clones, 172 Competition, 100, 106, 111, 118,
Clotting agents as venoms, 486 173, 621; defense mechanisms,
Clown fish, 624 125; distinguished from
Club mosses, 241 allelopathy, 15; and hierarchies,
Clutch size, 576; and clines, 81 329; interspecific, 644; and
Coal, Age of, 242 predation, 539; reduction
Cobras, 488 through migration, 409; and
Cocaine, 404 speciation, 545; and species
Codeine, 404 loss, 609; and territoriality, 634
Coenogonium leprieurii (lichen), 381 Competition exclusion law, 644
Coevolution, 10, 24, 86, 89, 174; Competitive exclusion, 66, 101
defense mechanisms, 127; Composting, 669
grasses and grazers, 304; Condensation, 338
pollinators, 495; and symbiosis, Condors, 203
624 Coniferous forests, 314, 629
Cohorts, 138 Conifers, 269, 629; evolution of,
Cold-blooded animals, 10 242
Collier, G., 115 Connectivity, 378
Colonies, ant, 343 Conservation biology, 35, 119, 433;
Colonization of the land, 90, 94 endangered plants, 209;
Coloration; agouti, 72; aposematic, wildlife management, 677. See
128, 167, 538; cryptic, 72; also Restoration ecology
marine animals, 396; warning, Conservation easements, 264
415 Consumers, 24, 52, 104, 174, 187,
Colubrids, 487 191, 557; macroscopic vs.
746
Subject Index
microscopic, 102. See also Craighead, John, 517

Heterotrophs Cranes, 203
Contact herbicides, 471 Crater Lake, 364
Contact zones, 275 Creep, 212
Continental shelf, 63 Cretaceous period, 243;
Continental slope, 63 coevolution during, 86
Continuous populations, 507 Cretaceous-Tertiary (KT) event,
Convention on International 244, 252
Trade in Endangered Species, Crickets, communication, 98
202 Crop plants; allelopathy, 15;
Convergence, 120, 122, 157, 615 diversity, 279; genetically
Cooperative behavior; altruism, modified, 284
19; and hierarchies, 330; mole Cross-dating, 146
rats, 389; social insects, 346 Cross-pollination, 495
Copepods, 393 Crossbills, competition among,
Copperhead snakes, 487 115
Copulation, 577 Crown fires, 260, 267
Coral reefs, 566, 624; territoriality Crustaceans, 393
in, 636 Cryoplanation, 655
Coral snakes, 487 Cryprochromes, 650
Corals, 566; Devonian period, 248; Crypsis, 72
parasites of, 625; reefs, 127; Cryptophytes, 588
stingers, 129 Cultural eutrophication, 222
Coriolis force, 154 Curare, 492
Corollas, 497 Cuvier, Georges, 237
Correlational analysis, 580 Cyanobacteria, 393, 465, 482
Correlational selection, 430 Cycads, evolution of, 242
Corridors, 378 Cycles of abundance, 513, 539
Costa’s hummingbirds, 158
Coterie (family unit), 388 Dance language of bees, 215, 345
Cottonmouth snakes, 487 Darwin, Charles, 13, 29, 38, 179,
Coumarins, 15 227, 237, 274, 428; description
Countershading, 72 of habituation, 320; ethology,
Countersinging, 98 215
Courtship; competition, 113; Darwin’s finches, 429. See also
displays, 167; isolating Finches
mechanisms, 361 Dasyatids, 488
Cowles, Henry Chandler, 181 Dating techniques;
Coyotes, 157, 300, 389, 665 dendrochronology, 146;
Crabs, hermit, 624 radiocarbon dating, 146
Craighead, Frank, 517 Davis, Michael, 323
747
Subject Index
Dawkins, Richard, 522 Department of the Interior, U.S.,

DDT, 256, 470, 473, 503; 200
dependence on, 351; species Deposition (water erosion), 211,
loss, 199; in zooplankton, 48 290
Death rates, 138; zoos, 438 Descent of Man and Selection in
Death rates (population analysis), Relation to Sex, The (Darwin),
137, 530 215, 238
Debt-for-nature swaps, 264 Descriptive ecology, 175
Deciduous forests, 314-315; Desert coyotes, 157
tropical, 586, 589 Desert Storm (military action),
Deciduous trees, 269, 629; 449
diseases, 41; savannas, 62; Desertification, 149, 153, 306
temperate forests, 59, 269 Deserts, 61, 149, 153-154, 160, 294,
Decomposers, 52, 104, 174, 187, 314; adaptations to, 9; cold, 315;
191, 256, 557, 561, 641; bacteria, hot, 315
290; lake bottoms, 369; Despotism, 330
zooplankton, 393 Detachment (water erosion), 211
Deep ecology, 123-124 Deterrent pheromones, 476
Deevey, Edward S., 139 Development; and ethology, 216;
Defense mechanisms, 102, 125, and evolution, 161, 166, 428;
130; and bioluminescence, 44; sustainable, 618, 620
plants, 88; and predation, 537; Devonian period, 93, 241
and territoriality, 634; venoms, Diagenesis, 370
486 Diamond, Jared, 33
Defensive associations, 128 Diatoms, 366, 482
Deforestation, 53, 131, 136, 150, Dichloro-diphenyl-trichloroethane.
263, 334; rain forests, 554 See DDT
Demes, 80, 274 Dieffenbachia (dumb cane plant),
Demographics, 137, 144. See also 491
entries under Population Differential growth, 650
Dendrochronology, 145, 148-149, Dimictic lakes, 367
464 Dinoflagellates, 482, 625
Density-dependent growth, 511 Dinosaurs, extinction, 249
Density-dependent migration, 410 Dioxin, 504
Density-dependent natural Directional selection, 429
selection, 579 Disasters, impact on communities
Density-dependent population and ecosystems, 106
fluctuations, 513, 577 Discrete displays, 96
Density-dependent territoriality, Discrete population, 507
637 Diseases; and biopesticides, 65;
Deoxyribonucleic acid. See DNA insect-borne, 41; urban and
748
Subject Index
suburban wildlife, 665; and Dusky seaside sparrows, 203

wildlife management, 678 Dust Bowl (1930’s), 213, 617
Dishabituation, 320 Dutch elm disease, 355
Dispersal, 38; corridors, 378; and Dwarfing, 62
landscape fragmentation, 375.
See also Seed dispersal Eagles; bald, 199; golden, 157
Dispersal of populations, 407 Earth Day (est. 1970), 181
Disphotic marine zone, 391 Earth First!, 123
Displays, 95, 113, 167, 170; and Earth Summit (1992), 312
territoriality, 634 Echinoderms, 397
Disruptive coloration, 72 Echolocation, 157
Disruptive selection, 429 Ecocentrism, 123
Disturbance in ecosystems, 189 Ecoenergetics. See Biomass;
Diterpenes, 404 Energy budgets; Energy flow;
Divergence, 120, 122 Food chain; Trophic levels;
Divergent populations, 12 Categorized Index
Diversity. See Biodiversity; Ecofeminism, 124
Biological phenomena Ecological isolation, 359
diversity; Ecosystem diversity; Ecological niches. See Niches;
Genetic diversity; Species Trophic levels
diversity Ecological pyramids, 186
DNA (deoxyribonucleic acid), 229 Ecological Society of America, 180
DNA sequencing, 233 Ecology; defined, 171, 178; history,
Dobzhansky, Theodosius, 358 179, 183
Dogs, social organization of, 389 Economy of Nature, The (Linnaeus),
Dominance hierarchies, 114, 329, 29, 179
637 Ecophysiology. See Categorized
Dominant species, 104 Index, Physiological ecology
Douglass, Andrew Ellicott, 145 Ecosystem diversity, 34
Dragonfish, 45 Ecosystem ecology; defined, 174.
Dragonflies, 158 See Biomes; Communities;
Drift. See Genetic drift Ecosystems; Categorized Index
Drought, 135, 150; dry tropics, Ecosystems, 32; agricultural, 352;
586; global warming, 294; coining of term, 255; and
grasslands, 62 communities, 100, 103;
Dry tropical biomes, 586 definition and history of the
Dual-process habituation- concept, 184, 190; and habitats,
sensitization theory, 320 314; invasive plants, 41; and
Dubautia (silversword), 14 pollution, 500; impact of
Duetting, 98 species loss, 197; studies, 191,
Dupes (mimicry), 415 195. See also Alpine tundra;
749
Subject Index
Arctic tundra; Biomes; Boreal Endangered Species Act (1990),

forests; Chapparal; Deciduous 454
tropical forests; Deserts; Endangered Species Preservation
Ecotones; Forests; Grasslands; Act (1966), 200, 610
Lake ecosystems; Marine Endemics (desert plants), 156
biomes; Mediterranean scrub; Endogenic chemical precipitates,
Mountain ecosystems; Old- 366
growth forests; Prairies; Rain Endomycorrhizae, 426
forests Endotherms, 172
Ecotones, 105, 631; tundra-forest, Endothia parasitica (fungus), 65
656; wetlands, 672 Energy budgets, 172, 174, 430;
Ecotoxicology. See carnivores vs. herbivores, 455;
Biomagnification; Pollution; and population size, 186; and
Pesticides; Categorized Index predation, 539
Ectomycorrhizae, 102, 426 Energy conversion, 642
Ectoparasites, 624 Energy expenditure and the food
Ectotherms, 172 chain, 256
Edge habitat, 375 Energy flow, 104, 177, 191; and
El Capitan reef complex, 568 biomass, 50, 54
El Niño, 591 English ivy, 41
Elapids, 487 Ephemerals, 157
Eldredge, Niles, 544 Epidemiology, 177
Elephant birds, 200 Epilimnion, 367
Elephant grass, 586 Epipelagic marine zone, 391
Elephant seals, territoriality, 637 Epiphytes, 102; endangered, 208
Elfin forest, 76 Episodic migrations, 410
Elicitors, 103 Epizoites, 622
Elm trees, 356 Equilibrium, and natural
Elton, Charles, 185, 255, 641 selection, 428
Elton’s pyramid, 185, 256 Equilibrium models of population
Embryology, 162 dynamics, 517
Emergent layer (rain forests), 552, Ericoid mycorrhizae, 426
558 Erosion, 211, 214; deserts, 62; fire-
Emigration, 508, 528 caused, 259; overgrazing, 306;
Empedocles, 236 reforestation, 574; slash-and-
Endangered species; animals, 196, burn agriculture, 591; soil, 150
204; and ecological niches, 647; Erosion control, 213
inbreeding of, 439; plants, 205, Essentialism, 237
210 Estrogens, environmental, 503
Endangered Species Act (1973), Estuaries, 100, 316, 391; algae in,
200, 503, 610 482, 592; tidal, 105
750
Subject Index
Ethograms, 219 Extinctions, 35, 109, 232, 246, 254,

Ethological isolation, 358 518, 608, 611; causes, 513; role
Ethology, 215, 221. See also of competition, 179; and
Behavioral ecology endangered species, 196;
Eucalyptus, 59, 271 caused by exotics, 177;
Euphorbs, 120, 492 inbreeding depression, 282; and
Eusocial colonies, 385 invasive species, 40; mass, 244,
Eutrophication, 222, 226, 483, 601; 246; passenger pigeons, 198;
cultural, 211; lakes, 225, 371. See plants, 241, 245; and pollution,
also Phytoplankton 500; and punctuated
Evaporation, 338 equilibrium, 544. See also
Evapotranspiration, 338 Species loss
Evenness of species, 33 Exxon Valdez oil spill, 446
Everglades (Florida), 206, 504
Evergreens, 59, 630. See also Falcons, 199
Coniferous forests; Conifers Fallow land, 590
Evolution, 227, 235; adaptations, Family units, mammalian. See
7; and altruism, 19; Mammalian social systems
coevolution, 86, 89; convergent, FAO. See United Nations Food
120, 122; and development, 161, and Agriculture Organization
166; divergent, 120, 122; history Fecundity, 140
of its study, 236, 240; isolating Feldspars, 594
mechanisms, 358, 363; natural Ferguson, Charles, 146
selection, 428; of plants, 241, Ferns, 241
245; puncutated equilibrium vs. Ferrets; black-footed, 203, 300; and
gradualism, 543, 548; and prairie dogs, 109
speciation, 605. See also Fertilization, failure, 360
Colonization of the land; Fertilizers, as soil contaminants,
Selection 601
Evolutionary ecology. See File, Sandra, 323
Evolution; Categorized Index Finches, adaptations of, 13. See
Evolutionary explosions, 246, 254 also Darwin’s finches
Exhaustible resources, 618 Fire ants, 345
Exotic species, 40, 110, 177, 648, Fire climax ecosystems, 258, 399
662; and species loss, 196 Fire management, 400
Experimental ecology, 175 Fire suppression, 260, 400
Explosions, evolutionary, 249 Fire worms, 45
Exponential population growth, Fireflies, 45, 96
531 Fires; chaparral, 78, 399; Devonian
Expression of the Emotions of Man period, 93; forest, 258, 262, 267,
and Animals (Darwin), 215 588, 631; in grasslands, 298;
751
Subject Index
nutrient loss from, 442; slash- ecosystems, 557; management,

and-burn agriculture, 591 263, 268, 422, 424; multiple use,
Firs, 629 422, 424; nutrient cycles in, 442;
Fish, 394; bioluminescent, 43; old-growth, 452, 454; rain
clown, 624; defense forests, 549, 553; temperate, 58-
mechanisms, 125; eutrophic 59; tropical, 59
environments, 224; herbivores, Fossil fuels, 501; global warming,
328; omnivorous, 456; 295; greenhouse effect, 310;
poisonous, 488; and pollution, tundra deposits, 657
504 Fossil record, 90, 232, 465, 546;
Fish and Wildlife Service, U.S., and theories of evolution, 250
196, 610, 677 Founder effects, 437
Fish lice, 622 Foxes, 300, 665; defense
Fisher, R. A., 522 mechanisms, 126; social
Fitness, 7; inclusive, 218; natural organization, 389
selection, 428; and reproductive Fragmentation, 374
strategies, 577 Franklin, Jerry, 452
Fitness variation, 281 Freshwater lakes, 315
Fixation, genetic, 282 Freshwater wetlands, 673
Flavonoids, 15 Fringing reefs, 564
Flavr Savr tomato, 284 Frisch, Karl von, 215, 345, 348
Flooding, 135; tolerance of, 672 Fritts, Harold, 147
Food chain, 47, 52, 104, 185, 255, Frogs; bullfrogs, 158; clines, 81;
257, 612; grasslands, 299; and poisonous, 489
herbivores, 328; and Fugitive species, 537
phytoplankton, 482 Functional response of predators,
Food pyramid (Elton’s), 641 537
Food webs, 53, 104, 174, 185, 255, Fundamentals of Ecology (Odum),
257, 612; and predation, 539 191
Foods, genetic modification of, Fungal associations, 425, 427
284, 287 Fungal diseases, 65, 355
Forbes, Stephen A., 29, 184, 189, 255 Fungi; as biopesticides, 66; lichens,
Forel, François A., 182 381; soil generation, 596
Forest fires, 258, 262, 267, 588, 631 Fungicides, 470
Forest Service, U.S., 265, 454, 677 Fungus gnats, 44
Forest zones, 551 Fur industry and species loss, 199
Forestry; multiple-use approach, Fynbos vegetation, 400
422, 424; sustainable, 572
Forests, 269, 273; boreal, 60, 629, Gaboon vipers, 488
632; coniferous, 314; deciduous, Gaia hypothesis, 30, 123
314; Douglas fir, 109; as Galápagos Islands, 121, 238
752
Subject Index
Game management, 677 Global ReLeaf program, 575

Game theory, 639 Global warming, 292, 297, 309,
Garstang, Walter, 163 504, 574; role of slash-and-burn
Geese; hierarchies, 331; agriculture, 591; and taiga, 631
imprinting, 216; snow, 435 Globigerina shells, 465
Gene flow, 80, 274, 277, 281; Gloger’s rule, 81
impact of corridors, 378; and Glowworms, 44
landscape fragmentation, 375 Glycosides, 491
Gene pools, 274, 576; defined, 229; Golden eagles, 157
and habitats, 313 Goldschmidt, Richard, 165
Genes, 274; regulatory, 164 Golley, Frank B., 192
Genetic diversity, 35, 278 Gondwanaland, 247
Genetic drift, 281, 283, 428, 435- Goodall, Jane, 324
436, 439, 522, 604; and Goosefoot family, 157
population fluctuations, 518 Gould, Stephen Jay, 163, 544
Genetic engineering, 433. See also Governing Nature (Linnaeus), 29
Genetic modification Graded displays, 96
Genetic hitchhiking, 281 Gradualism, 231, 543, 548
Genetic modification, food crops, Grand Canyon National Park and
284, 287 Game Preserve, Arizona, 301
Genetic pollution, 286 Grass, elephant, 586
Genetics; behavioral, 217; and Grass frogs, 81
evolution, 228; Mendelian, 162, Grasslands, 57, 62, 149, 298, 303,
239; populations, 520, 527 314
Genotypes, 521 Grasslands management, 301
Geochemical cycles, 288, 291. See Grasslands Project, 192
also Carbon cycle; Hydrologic Graunt, John, 28
cycle; Nitrogen Cycle; Nutrient Gravitropism, 651
cycles; Oxygen cycle; Grazing, 304, 307, 560; forests, 265;
Phosphorus cycle effect on grasses, 304; symbiotic
Geoffroyism, 237 characteristics, 299
Geographic speciation, 604 Great Barrier Reef, 564
Gestation, and reproductive Great Basin Desert, 156
strategies, 576 Great Lakes, North America, 504
Gila monsters, 486 Greenhouse effect, 54, 292, 308,
Gill slits, 162 312; rain forests, 557
Glaciers, 295, 310 Greenhouse gases, 152, 292, 671;
Glassfish, 623 and deforestation, 135
Gleason, Henry A., 30, 182 Grinnel, Joseph, 641
Global ecology. See Categorized Grooming, 95
Index Gross production, 51, 172
753
Subject Index
Ground fires, 268 Hale, Matthew, 28

Groundwater, 338, 668 Hamilton, William D., 349
Groundwater pollution, 668 Hanford Nuclear Reservation, 670
Group dynamics, 18 Haplodiploidy, 344
Group selection, 431 Hardin, Garrett, 302
Groups; and cooperation, 346; Hardwoods, 270
urban and suburban wildlife, Hardy-Weinberg law, 435, 521, 525
661 Harrison, Benjamin, 422
Grouse, competition among, 114 Hartig nets, 427
Growth, differential, 650 Harvesting and nutrient loss, 442
Growth rates (population Hatchetfish, 45
analysis), 530 Haustoria, 382
Growth rings. See Hawaiian silversword alliance, 13
Dendrochronology Hawks, 665
Guam rails, 203 Hazardous waste, 667
Gully erosion, 212 Heat islands, 296
Gymnosperms, evolution of, 241 Heavy metals, 48
Gypsy moths, 41, 65 Hedera helix (English ivy), 41
Heliotropism, 653
Habitat destruction. See Habitat Hemotoxins, 486
loss Herbicides, 285, 470-471; invasive
Habitat isolation, 359 plant control, 355
Habitat loss, 177, 196, 376, 647; Herbivores, 102, 111, 174, 256, 304,
deserts, 159; environmental 326, 328, 455, 641; as parasites,
change, 252; Madagascar, 200; 174; as pollinators, 86;
mountains, 420; plant species, rangeland animals, 561
205; through pollution, 500; and Herbivory, 490
population fluctuations, 518; Herd behavior, 388
rain forests, 552; wetlands, 675; Heredity, laws of, 228
and wildlife management, 678 Hermit crabs, 624; defense
Habitats, 313, 318, 659; mechanisms, 125
demographics of, 137; and Heterobasidion annosum (pine
ethology, 216; landscape pathogen), 67
ecology, 374, 380; management, Heterochrony, 162
677; marine, 391; protective, Heterogeneity. See Biodiversity
127; and territoriality, 633 Heterotrophs, 52, 86, 111. See also
Habituation, 319, 325 Consumers
Hadalpelagic marine zone, 392 Heterozygote genotypes, 435
Haeckel, Ernst, 161, 179 HFCs. See Hydrofluorocarbons
Hairston, Nelson G., 255 Hibernation, 9, 412; desert
Haldane, J. B. S., 522 animals, 158
754
Subject Index
Hierarchies, 19, 329, 332; and Hydrosphere, 69

territoriality, 637 Hydrotropism, 653
High-grading, 132 Hymenoptera (ants, bees, wasps),
History of ecology, 171, 178; 19, 343
ecosystems, 184, 190 Hyperarid climates, 150
History of evolutionary theory, Hyperresponsiveness, 321
227, 235 Hypolimnion, 368
Hitchhiking, genetic, 281 Hypoxia in eutrophic
Homeostasis, 346 environments, 223
Homeotherms, 10
Homing pigeons, 407 IBP. See International Biological
Homozygotes, 522 Program
Honest signaling, 167 Ice ages, 295, 310
Hooke, Robert, 28 Ice-rafting, 366
Hookworms, 625 Iguanas, 158
Hormones; auxins, 650; Immigration, 508, 528
pheromones, 476, 481 Immutability of species, 236
Horses, 301 Imprinting, 216
Horsetails, 241 Inbreeding, 274, 436, 522, 605
Host species, 622 Inbreeding depression, 282, 495
Hubbard Brook project, 188, 193, Incineration, 669
442 Inclusive fitness, 218
Human ecological impacts, 176 Indicator species, 155
Human population growth, 257, Industrial Revolution, 1
272, 333, 337, 375; fire Infiltration capacity of soil, 340
suppression, 401 Inhibition and habituation, 322
Humboldt, Alexander von, 181 Innovation and evolutionary
Hummingbirds, Costa’s, 158 explosions, 252
Humphrey, G., 322 Inquilinism, 623
Hunting, species loss, 196; Insect communication, 345
predation, 389, 539. See also Insect societies, 95, 343, 350
Poaching Insecticides, 470
Hurricane Mitch (1998), 592 Insects; as biopesticides, 67;
Huxley, J. S., 165 herbivorous, 326; invasive, 41;
Hybrid sterility, 360 omnivorous, 456; pheromones,
Hybrid swarms, 83, 359 478; poisonous, 487;
Hybrid zones, 80, 85, 275, 604 pollinators, 496; stink bugs, 126;
Hybridization, 275 symbioses with plants, 326
Hydrofluorocarbons, 308 Instinctive behavior, 217
Hydrologic cycle, 69, 338, 342 Integrated pest management, 351
Hydrophytes, 672 Interbreeding, 358, 543, 604
755
Subject Index
Interception (hydrologic cycle), IUCN. See World Conservation

339 Union
Intergenerational justice, 618 Ixtoc I, 446
Internal nutrient cycling, 440
International Biological Program, Jackrabbits, 158
180, 192 Jacobson’s organ, and
International Council for Bird pheromones, 478
Preservation, 196 Jellyfish, 45; defense mechanisms,
International Panel on Climate 130
Change, 311 Johnson, Roswell, 641
International Union for Jurassic period, 243
Conservation of Nature and Juvenile survivorship patterns,
Natural Resources. See World 139
Conservation Union
Interspecific competition, 113 K strategy, 81, 578
Intersterility, 360 K/T boundary. See Cretaceous-
Intertidal marine zone, 63, 391 Tertiary (KT) event
Intraspecific competition, 113, Kaibab Plateau (Arizona) deer
645 disaster, 301
Intrinsic rate of increase, 530 Kairomones, 477
Introduced and exotic species, Kandell, E. R., 321, 323
583 Kaufman, Thomas, 163
Introgression, 80, 85, 275 Kestrels, American, 665
Introgressive demes, 83 Keystone predators, 106
Introgressive hybridization, 275 Keystone species, 104, 647; and
Invasive plants, 354, 357 predation, 539
Invasive species. See Biological Kidneys of desert animals, 159
invasions; Invasive plants Killer algae, 357
IPCC. See International Panel on Kimura, Motoo, 523
Climate Change Kin selection, 20, 218, 344, 431,
IPM. See Integrated pest 522
management King Solomon’s Ring (Lorenz), 216
Iridium, as a cause of extinction, Krebs, Charles, 111
247 Krummholz region, 656
Island biogeography, 33, 38 Kudzu, 41
Isoflavones, 405 Kyoto Accords (1997), 312
Isolating mechanisms, 83, 358,
363; mountains, 419 “Lake as a Microcosm, The”
Isoptera (termites), 343 (Forbes), 29
Isotherms, 61 Lake ecosystems, 185, 315, 364,
Iteroparous species, 579 373; pollution effects, 189
756
Subject Index
Lake Tahoe, eutrophication in, 222 Lizards, 486; black-collared, 158

LaMarche, Valmore, 146 Locusts, population fluctuations,
Lamarck, Jean-Baptiste, 29, 179, 515
227, 236 Logging; deforestation, 132;
Land management, 422, 424 national forests, 265; old-
Landfills, 668 growth forests, 453;
Landscape ecology, 374, 380 reforestation, 572
Landscape fragmentation, 374 Logistic population growth, 531,
Lantern fish, 45 577
Laterization, 572 Longevity, 142
Layering, 630 Lorenz, Konrad, 215
Leaching, 441, 601 Lotic communities, 315
Leafy spurge, 356 Lotka, A. J., 115
Learning; and ethology, 216; Lotka-Volterra model, 541
habituation, 319 Love Canal, 670
Lechuguilla, 156 Lovelock, James, 30
Leeuwenhoek, Antoni van, 28 Luciferin and luciferase, 43
Leks, 114, 167 Lucretius, 236
Lemmings, 514 Lycopene, 405
Lemurs, 200, 552 Lycophytes, 241
Lentic communities, 315 Lyell, Charles, 29, 179, 238
Leonardo da Vinci, 145 Lynxes, 514
Leopard frogs, 82 Lysimeters, 597
Leopold, Aldo, 677
Lice, 622 Macaque hierarchies, 331
Lichens, 381, 384; soil generation, MacArthur, Robert H., 38, 115,
596 578
Life cycles, 513 Macrohabitats, 313
Life spans, 138; urban and Macrotermes natalensis (termites),
suburban wildlife, 661 347
Life tables, 143 Madagascar, species loss on, 200
Lignin, 405 Mallee vegetation, 400
Likens, Gene E., 193 Mammalian social systems, 385,
Limnology, 182, 364, 373 390; hierarchies, 332
Lindeman, Raymond L., 186, Mammals, omnivorous, 456
191 Mangroves, 674
Linnaeus, Carolus, 29, 179, 236 Man-o’-war, 393
Lions, competition among, 114 Maquis vegetation, 400
Lithosphere, 69 Marbled murrelets, 109
Litter size, 576 Marcus, Emilie A., 322
Littoral lake zone, 371 Margulis, Lynn, 30
757
Subject Index
Marine biomes, 316, 391, 398; Metabolism, desert animals, 159

phytoplankton, 483. See also Metabolites, 25, 126-127, 402, 406;
Ocean biomes coevolution of, 88; secondary,
Marine ecology, 182 490. See also Poisonous animals;
Marine regression and extinctions, Poisonous plants
246 Methane, 668; and global
Marine zones, 391 warming, 295, 310
Mark-recapture methods, 109, 116, Methuselah Tree, 146
516, 533 Mice, 665
Marker pheromones, 476 Microcosms, 185
Marshes, 391, 674 Microhabitats, 313
Mathematical ecology, 176 Migration, 12, 274, 407, 414, 428;
Mathematical models; and natural and hybrid swarms, 83;
selection, 432; population invasive species, 40;
genetics, 524; population mammalian social
growth, 529; predation, 541; organization, 388; and
predator-prey relationships, predation, 537
513 Migrations, birds, 514
Mating; calls, 359; hybrids, 83; Milkweed plant, 538
pheromones, 477; systems, 577. Milpa agriculture. See Slash-and-
See also Reproduction burn agriculture
Matorral vegetation, 400 Mimicry, 25, 73, 97, 128, 415, 418;
Maupertuis, Pierre-Louis Moreau aggressive, 538; Batesian, 128,
de, 236 538; fireflies, 45; Müllerian, 128,
Maximal intrinsic rate of increase, 538
530 Mineralization, 441
Mayr, Ernst, 543 Miocene epoch, 244
Mech, David, 517 Missing links, 236
Mechanical isolation, 359 Mistletoe, 102
Meckel, J. F., 161 Möbius, Karl, 182
Medicinal plants; alkaloids, 404; Models (mimicry), 415
endangered, 209 Mollusks, 397
Mediterranean scrub, 62, 399, 401 Monimolimnion, 372
Mendel, Gregor, 229, 239 Monkeys; communication, 98;
Mendelian genetics, 521 hierarchies, 331; social systems,
Mercury poisoning, 48 386
Mesopelagic marine zone, 391 Monoclimax theory, 615
Mesotrophic lakes, 225 Monoculture, 264, 306
Mesozoic era, 243 Monogamy, 577
Metabolic adaptations, 9 Monotropoid mycorrhizae, 426
Metabolic pathways, 402 Morphine, 404
758
Subject Index
Mortality rates. See Death rates National Park Service, U.S., 677
Mother-young relationships, 385 National Wildlife Federation,
Moths; peppered, 428; 678
pheromones, 478; as National Wildlife Refuge System,
pollinators, 497 200
Mount Tamalpais, California, 78 Native plants, 40
Mountain ecosystems, 419, 421 Natural selection. See Selection;
Mountain lions, 385 Selection regimes; Sexual
Mousse (oil spill), 444 selection
Müllerian mimicry, 128, 416, 538 Nature preserves, 33. See also
Multiple-use approach, 422, 424 Parks; Wildlife refuges; Zoos
Multiple Use-Sustained Yield Act Navigation, 407
(1960), 423 Nectar, 496
Murrelets, marbled, 109 Negative assortative mating, 435
Musk, 127 Nekton, 393
Mutagens, 438 Nematocides, 470
Mutational meltdown, 283 Nematocysts, 129
Mutations, 229, 274, 428, 435, 439, Neoteny, 163
520; and divergence, 12; and Neritic marine zone, 63, 391
natural selection, 7; random, Nested hierarchies, 330
239 Nesting; social insects, 346;
Mutualisms, 24, 87, 100; and territoriality, 635
coevolution, 174; pollinators, Net production, 51, 172
495; distinguished from Neurobiology, 216
symbiosis, 623. See also Neurotoxins, 486
Symbioses Neurotransmitters and
Mycobionts, 381, 425 habituation, 321
Mycorrhizae, 101, 425, 427; in old- Neutral mutation, 524
growth forests, 453 Neutralisms, 100
New forestry, 268
Naess, Arne, 123 Niche generalizations, 645
NASA. See National Aeronautics Niche overlap, 644
and Space Administration Niches, 13, 419, 641, 649; and
Natality rate, 138 competition, 113; competition
National Aeronautics and Space for, 115; defined, 171; and
Administration, 461 hybridization, 83
National Environmental Nicotine, 404
Protection Act (1970), 677 Nightshade family, 404
National Oceanic and Nimbus meteorological satellites,
Atmospheric Administration, 461
449, 460 NIMBY (not in my backyard), 669
759
Subject Index
Nitric oxides, 501 Offspring care; altruism, 19-20;

Nitrogen, excessive, 222 camouflage, 74; territoriality,
Nitrogen cycle, 185, 289, 441-442 635
“No net loss” policy, 584 Oil deposits (tundra), 657
NOAA. See National Oceanic and Oil spills, 444, 451
Atmospheric Administration Old-growth forests, 133, 265, 452,
Nocturnal animals, 26, 158, 315, 454
359, 488, 663 Olfactory displays, 169
Nolen, Thomas G., 322 Oligotrophic lakes, 225
Nomadic animals, 407 Omnivores, 112, 257, 455-456
Nonbranching evolution, 228 On the Origin of Species by Means of
Nongeographic speciation, 605 Natural Selection (Darwin), 29,
Nonrandom mating, 435, 439 179, 215, 238, 274
Nonsymbiotic mutualism, 25 Ontogeny, 161
North American biomes, 314 Operation Desert Storm, 449
North American prairies, 299 Operators (mimicry), 415
North American taiga, 629, 632 Opossums, 665
North American Wetlands Optimality theory, 639
Conservation Act (1989), 675 Opuntia (prickly pear cactus), 120
North American Wetlands Orchidaceous mycorrhizae, 426
Conservation Reauthorization Orchids; endangered, 208;
Act (2002), 675 pollination, 498
Noss, Reed F., 34 Ordovician period, 90
Nuclear and radioactive waste, Organismal ecology, defined, 171
669 Organophosphates, 471
Nuclear weapons, 669 Orians, G. H., 115
Numerical response of predators, Orientation response, 321
536 Ornithischians, 158
Nutrient cycles, 185, 440, 443; soil Outcrossing, 286
generation, 596. See also Overcultivation, 150
Geochemical cycles Overgrazing, 150, 300, 304, 307,
Nutrients, excess, 222, 226 336, 561; southern Africa, 207
Overland flow, 340
Oak Ridge National Laboratory, Ovulation, 577
180, 188 Owls; barn, 665; screech, 665;
Ocean biomes, 63, 391. See also short-eared, 662; snowy, 662;
Marine biomes spotted, 109, 203, 453
Ocean pollution, 444, 451 Oxygen cycle, 288; rain forests,
Oceanic marine zone, 391 555
Octopuses, mimicry in, 417 Oxygen depletion, eutrophic
Odum, Eugene P., 191 environments, 223
760
Subject Index
Ozone, 556 Pelagic marine zone, 391

Ozone depletion, 310, 457, 463, Pelicans, 203
501 Peniophora gigantea (biopesticide),
Ozone hole. See Ozone depletion 67
Peppered moth, 428
Pack structure (dogs), 389 Peppered moth adaptations, 10
Paclitaxel, 404 Per capita rate of growth, 530
Pair-bonding displays, 168 Per capita rate of increase, 509-510
Paleobotany, evolutionary Per capita rates, 509
theories, 543, 548 Perception, variations among
Paleodiversity, 251 organisms, 215
Paleoecology, 37, 464, 469; Peregrine falcons, 199
methods, 188 Pereskia (cactus), 120
Paleogene-Neogene period, 244 Perfluourocarbons, 308
Paleontology, 232, 543 Permafrost, 61, 629, 655
Paleozoic era, 241 Permian-Triassic extinction, 242,
Paley, William, 237 252
Palos Verdes butterflies, 203 Persian Gulf, 450
Pampas, South American, 298 Pest control; pheromones, 478;
Parapatric speciation, 604 and pollution, 503
Parasites; cowbird, 377; edge Pesticides, 470, 475;
species, 377; and predation, 536 biomagnification of, 47;
; zooplankton, 393 biopesticides, 65, 68; integrated
Parasitism, 87, 125, 174, 621, 624 pest management, 351;
Parks, 379. See also Nature resistance to, 351; species loss,
preserves; Wildlife refuges; 199. See also DDT
Zoos Petroleum, origins, 234
Particulate pollutants, 591 PFCs. See Perfluourocarbons
Passenger pigeons, extinction of, Phenolics, 15, 405
198 Phenotypes, 435
Patch reefs, 565 Pheromones, 95, 170, 220, 476, 481;
Patches of habitat, 375 mating behavior, 359; social
Pathogens, 102 insects, 345
Pavlov, Ivan, 322 Philodendrons, 491
PCBs, 48, 504, 601 Philosophical Account of the Works of
Pearl, Raymond, 139 Nature, A (Bradley), 28
Pearlfish, 623; sea cucumber Phoresis, 622
associations, 129 Phosphate runoff, 225
Peat, 242 Phosphates, 222
Pecking orders, 113, 331 Phosphorus, excessive, 222
Pelagial lake zone, 371 Phosphorus cycle, 290
761
Subject Index
Photic marine zone, 391, 483 Point-quarter technique, 117

Photinus (firefly), 45 Poison ivy, 493
Photobionts, 381; in lichens, 382 Poison oak, 493
Photodissociation, 555 Poison sumac, 493
Photophores, 43 Poisonous animals, 486, 489
Photosynthesis, 50, 69, 104; CAM Poisonous plants, 490, 494
and C4 , 156; as an oxygen Polar bears, 13
source, 556; as primary Pollen, 367, 495-496; ancient, 464
productivity, 187 Pollination, 10, 101, 495, 499
Phototropism, 650 Pollinators, 26; coevolution of, 86
Photuris (firefly), 45 Pollinia, 498
Phylogeny, 161 Pollution; and ecosystems, 189;
Physiological adaptations, 9 effects, 500, 506; eutrophication,
Physiological ecology. See 222, 226; oceans, 444, 451; soil,
Categorized Index 601, 603; and urban and
Phytoalexins, 103 suburban wildlife, 659; and
Phytochemicals, 490 wildlife management, 678
Phytoecdysones, 405 Polychaete worms, 397
Phytoplankton, 392, 482, 485; Polychlorinated biphenyls. See
eutrophic environments, 223. PCBs
See also Eutrophication Polyclimax theory, 615
Pianka, Eric, 578 Polyculture, 264
Pied wagtails, 636 Polygamy, 577
Pigeons; migration, 407; Polyploidy, 605
passenger, 198; urban, 661 Polyterpenes, 405
Pigments, 405 Population analysis, 117, 507, 512;
Pinchot, Gifford, 422 demographics, 137; and
Pine family, 629 wildlife management, 678
Pinnipeds, 396 Population ecology, 172, 317. See
Plankton, 63, 392; migrations, 409. also Population analysis;
See also Phytoplankton; Categorized Index
Zooplankton Population fluctuations, 513, 519;
Plant-animal interactions. See and pollution, 501; and
Animal-plant interactions predation, 539
Plant ecology, 181 Population genetics, 435, 520,
Plant Quarantine Act (1912), 356 527
Plant Succession (Clements), 30 Population growth, 138, 172, 528,
Plants, land vs. sea, 91 535, 577; human, 333, 337
Plate tectonics, 419 Population regulation, 173, 176
Poaching, 678. See also Population size, and genetic drift,
Endangered species; Hunting 436
762
Subject Index
Populations; and habitats, 313; Production biology, 256

human, 257, 272, 401 Productivity (rate of accumulation
Porcupine defense mechanisms, 125 of biomass), 50, 187
Positive assortative mating, 435 Profundal lake zone, 371
Post-extinction recoveries, 249 Programmed migrations, 409
Postmating isolation, 359 Progymnosperms, 242
Prairie dogs, 300; and ferrets, 109; Prometheus Tree, 147
social organization, 388 Protective species, 128
Prairie potholes, 673 Protozoa as biopesticides, 67
Prairies, 298, 303; North Psittocosis, 665
American, 298, 315 Psychology, comparative, 216
Precipitation, 339; global Pterophyta (ferns), 241
warming, 294 Pueraria lobata (kudzu), 41
Predation, 88, 106, 125, 536, 542; Puff adders, 488
balance of nature concept, 255; Punctuated equilibrium, 231, 543,
and bioluminenscence, 44; and 548
camouflage, 74; mammalian Purple loosestrife, 356
social systems, 387; marine Pyramid, Elton’s, 185, 256
animals, 395; mimicry in, 415; Pyrethrins, 67
relative rarity, 186; and urban
and suburban wildlife, 664; Quadrat method, 516
venoms, 486; wasps, 344. See Quaternary period, 245
also Predator-prey relationships Quiet Crisis, The (Udall), 422
Predator-prey relationships, 25, Quills, 125
174, 609, 621; and population Quinine, 404
fluctuations, 514. See also Quinone, 127
Predation
Premating isolation, 358 r strategy, 81, 578
Prescribed burns, 260 Rabbits; Australian, 431; as
Preserves, 379 exotics, 110; hierarchies, 331
Prickly pear cactus, 120, 156 Raccoons, 665; defense
Primary producers. See Producers mechanisms, 126
Primary productivity. See Radioactive fallout, 188
Productivity Radiocarbon dating, 146
Primary succession, 613 Raff, Rudolf, 163
Primates; behavior, 216; social Rails, 203
systems, 386 Rain forests, 57, 59, 263, 549, 553;
Primer pheromones, 95, 476 atmospheric interactions, 177,
Producers, 24, 50, 104, 111, 174, 554, 559; Cenozoic, 244;
187, 191, 482, 557, 561, 641. See destruction rates, 336;
also Autotrophs ecosystems, 557; evolution of,
763
Subject Index
242; slash-and-burn agriculture, patterns, 140; pheromones, 478;

590; subtropical, 314; and population growth, 529;
temperate, 58, 549, 557, 629; sexual, 604; strategies, 171, 435,
tropical, 52, 55, 102, 131, 203, 439, 576, 582; and terrioriality,
263, 269, 549, 557, 574, 615. See 636-637
also Tropics Reservoirs, 365
Rain-shadow effect, 154 Residence time (hydrologic cycle),
Rainbow trout, 313 341
Rangeland, 560, 563 Resilience in ecosystems, 108
Ranges of species, 313 Resistance, predicted by
Rank distinctions, 330 population genetics, 526
Rankin, Catherine H., 322 Resistance to ecosystem
Rats, 665; social organization, 385 disturbance, 108
Rattlesnakes, 158, 487 Resistance to pesticides, 472
Ravens, 157 Resources, exhaustible, 618
Ray, John, 28 Respiration, 70, 172, 187, 441, 554
Recapitulation, 161 Restoration ecology, 583, 585;
Reciprocal sacrifice, 20 endangered species, 203;
Reciprocal signaling, 167 eutrophicatication, 225; tundra
Recombinant DNA technology, biomes, 657. See also
284 Categorized Index
Recycling, 175, 668 Retardation, 163
Red Data Book (IUCN), 202 Reticular network (vertebrates),
Red-tailed hawks, 663 321
Red tides, 224, 483. See also Algal Rhesus monkey hierarchies, 331
blooms; Brown tides; Rhyniophytes (early land plants),
Eutrophication; Phytoplankton 92, 241
Red tree voles, 109 Ricin, 492
Red wolves, 116 Rill erosion, 211
Reefs, 564, 571; protective Riparian (river) ecosystems, 301
habitats, 127 River ecosystems. See Riparian
Reforestation, 264, 572, 575 ecosystems
Regeneration; defense rmax , 530
mechanisms, 129 Rocky Mountain National Park,
Releaser pheromones, 476 Colorado, 420
Remoras, 623 Rodenticides, 470
Repellant pheromones, 476 Rodents; behavior, 216; as
Reproduction; asexual, 282, 524, pollinators, 497; social
605; demography of, 140; organizations, 388
isolating mechanims, 358, 363; Roosevelt, Theodore, 423
lichens, 383; migration, 409; Rubber, 405
764
Subject Index
Rubber trees, 552 Seasonal isolation, 359

Rudist reefs, 568 Secondary metabolites, 25, 127
Ruminants, 305, 327, 455 Secondary succession, 613
Runoff; agricultural, 213; as a Sedimentation; lakes, 211, 365;
pollutant, 601 storm drains, 259
Rusts, white pine blister, 41 Seed dispersal, coevolution of, 88
Seed germination and fire, 258,
Sacrifice, reciprocal, 20 400
Sagebrush, 156 Seedless vascular plants, 241
Saguaro cactus, 156 Selection (natural), 7, 12, 171, 217,
Sahara Desert, 156, 207 227, 238, 344, 428, 434, 521, 576;
Sahel Desert, 150 and altruism, 18; defense
Saint-Hilaire, Étienne Geoffroy, mechanisms, 125; and demes,
237 80; disruptive, 605; genetic
Salicylic acid, 405 drift, 281; isolating
Salt marshes, 674 mechanisms, 358; sexual, 238,
Saltation (wind erosion), 212 430; species selection vs., 545
Saltwater wetlands, 673 Selection regimes, 514
Sampling effect, 436 Selective harvesting, 132
Santa Ana winds, 399 Selective sweeps, 282
Saprotrophs, 112 Self-pollination, 495
Saurischians, 158 Selfish genes, 522
Savannas, 58, 62, 298, 586, 589 Semelparous species, 579
Scale of being, 161, 236 Semiarid climates, 149
Scavengers, 344, 455, 641; urban, Semigeographic speciation, 604
661 Semiochemicals, 476
Schistosomes, 625 Sense organs and camouflage, 75
Schulman, Edmund, 146 Sensitization, 319, 325
Sclerophyllous forests, 587 Sequoias, 146
Scorpaenidae (venemous fish), 489 Seres, 612
Scorpions, 158, 487 Serotonin, 322
Scrub. See Mediterranean scrub Serres, E. R., 161
Sea anemones, 624 Serres-Meckel law, 162
Sea cucumbers, evisceration, 129 Sex and territoriality, 636
Sea hares, 321 Sex ratios, 141, 508; population
Sea levels, rising, 295 analysis, 511
Sea turtle migration, 408 Sexual reproduction, 172, 604
Sea urchins, 126 Sexual selection, 238, 430
Seabirds, 394 Sheep, bighorn, 157
Seals; elephant, 637; overhunting Sheet erosion, 211
of, 199 Sheet flow, 341
765
Subject Index
Shells, as defense mechanisms, Society of American Foresters, 268

125 Sociobiology, 218
Shells, sea, 465 Softwoods, 270
Sherrington, Charles, 323 Soil, 594, 600; contamination, 601,
Sickle-cell disease, 520 603; degradation, 572; erosion,
Sierra Club, 123, 678 211, 214, 591; nutrients in, 440;
Sierra Nevada, 77 sampling, 597; sterilants, 471; in
Signal pheromones, 476 waste management, 600;
Silent Spring (Carson), 30, 193, 200 wetlands, 672. See also Erosion;
Silurian period, 90, 92 Soil chemistry
Silverswords, 13 Soil chemistry, 597
Similarity (for community Soil conservation, 213
classification), 109 Soil Conservation Service, 677
Single larger or several smaller Soil contamination, 131
(SLOSS) controversy, 33, 379 Sokolov, E. N., 320, 322
Siphonophores, 45 Solar radiation, 69; greenhouse
Skunks, 665; defense mechanisms, effect, 308
127 Solar tracking, 653
Slash-and-burn agriculture, 264, Solenopsis saevissima (fire ants), 345
272, 590, 593; and deforestation, Solid waste, 667
132; global warming, 295; Sonoran Desert, 156
reforestation, 572 Source reduction, 670
Sleep, 321 South American dry forests, 588
Slobodkin, Lawrence B., 255 Sparrows, dusky seaside, 203
SLOSS (single larger or several Speciation, 239, 276, 604, 607;
smaller) controversy, 33, 379 adaptive radiation, 12, 14; and
Sludge treatment and disposal, isolating mechanisms, 362;
669 punctuated equilibrium, 543,
Smell; displays, 169; and 548. See also Categorized Index
pheromones, 478 Species; definitions, 228;
Smith, Frederick E., 255 formation, 543; population
Snags, 452 fluctuations, 513; and
Snail darters, 202 territoriality, 633
Snakes, venomous, 487 Species diversity, 32
Snow geese, 435 Species loss, 376, 608, 611;
Snowpack, 341 animals, 196, 204; plants, 205,
Snowshoe hares, 514 210. See also Extinctions
Social Darwinism, 234 Species-removal studies, 609
Social ecology, 124 Species selection, 545
Social systems; insects, 95, 343, Sphenophyta (horsetails), 241
350; mammals, 385, 390 Sphinctozoans, 568
766
Subject Index
Spiders, 487 Sudbury, Ontario, emissions, 1

Spines (defense mechanisms), 126, Sugars, 50
489 Sulfur dioxide, 501
Splash erosion, 211 Summerwood, 145
Spotted owls, 109, 203, 453 Sunflower family, 157
Springwood, 145 Superfund, 670
Spruce family, 629 Supreme Court, U.S., 202
Spurge family, 120 Surface fires, 267
Squid, bioluminescence in, 43 Survivorship patterns, 138
Squirrels, 158 Suspension (wind erosion), 213
Stability thresholds, 110 Sustainable agriculture, 593
Stabilizing selection, 429 Sustainable development, 618, 620
Stagnation, 222 Sustainable forestry, 264-265, 572
Stanley, Steven, 545 Sustained yield, 422
Starling, European, 662 Swidden agriculture. See Slash-
Startle response, 319 and-burn agriculture
Status signs, 167, 331 Swifts, Vaux’s, 109
Steppes; Eurasian, 298; short- Switching, 537
grass, 57 Symbioses, 24, 87, 621, 628; and
Steroids, 405 bioluminescence, 43; lichens,
Stimulant pheromones, 476 381; mycorrhizal, 425, 427. See
Stimulus; generalization, 320; also Mutualisms
response, 319 Sympatric speciation, 605
Stingrays, 488; defense Symphilism, 623
mechanisms, 126 Systemic herbicides, 471
Stings, 129
Stink bugs, 126 Tactile communication, 99
Stonefish, 489 Tactile displays, 169
Stratigraphy, 467 Taiga, 629, 632, 656
Stromatolites, 465, 567 Tall-grass prairies, 560
Stromatoporoids, 568 Tannins, 15
Structural adaptations, 9 Tansley, Arthur G., 186, 255
Strychnine, 493 Tapeworms, 622
Study of Instinct, The (Tinbergen), Taxus (yew), 492
216 Teakwood, 552
Substrate preferences of Tegitecula-yucca interaction, 24
organisms, 316 Tellico Dam, Tennessee, 202
Succession, 105, 173, 583, 612, 617; Temperate forests, 58-59, 269, 558
wildlife management, 678 Temperature of Earth. See Global
Succulents; adaptations, 156; warming
evolution of, 120 Temporal isolation, 359
767
Subject Index
Tendrils, 652 Transparency, 74

Tennessee Valley Authority, 202 Transpiration, 338; and
Termite-protozoan mutualisms, deforestation, 135
174 Transplant experiments, 116
Termites, 344; African, 347 Transport (water erosion), 211
Terns, arctic, 411 Trebouxia (algae), 381
Terpenes, 15 Tree farming, 572
Terpenoids, 404 Tree-of-heaven, 659
Terracing, 213 Tree rings. See Dendrochronology
Terrestrial communities, 173 Trees; endangered, 552; and global
Territoriality, 112, 633, 640; warming, 574
mammalian social Triassic period, 243
organization, 388; mice, 117 Trichoderma (biopesticide), 67
Tetraterpenoids, 405 Trimerophytophyta (early land
Thallus, lichens, 382 plants), 241
Thanatocoenoses, 466 Triterpenoids, 405
Theophrastus, 15 Trophallaxis, 99, 345
Theory of Island Biogeography, The Trophic levels, 104, 174, 187, 256,
(MacArthur and Wilson), 578 562, 641, 649; and predation,
Thermocline, 368 539
Thermoregulation in deserts, 158 Tropical forests, 57, 59, 586, 589
Thienemann, August, 256 Tropics, habitat loss, 177. See also
Thigmotropism, 652 Rain forests
Threat displays, 167 Tropisms, 650, 654
Threatened species, 196 Trypanosomes, 625
Tillage practices, 213 Tundra, 56, 61, 314, 631, 655, 658
Tillandsia (epiphyte), 102 Turbidite deposits, 366
Timber industry, 132; national Turkeys, 158
forests, 265; old-growth forests,
452; reforestation, 572 Udall, Stewart, 422
Tinbergen, Nikolaas, 215 Ultraviolet radiation, 457, 462,
Toads, 158; poisonous, 489 501
Topsoil, 211, 214; erosion, 336, Umbilicaria esculenta (lichen), 383
574 Understory (forests), 269, 551, 558
Toxic chemicals, 47 Ungulate social organization, 388
Toxicodendron radicans (poison Uniformitarianism, 238
ivy), 493 United Nations Environmental
Trace fossils, 467 Program, 462
Trail pheromones, 476 United Nations Food and
Transeau, Nelson, 256 Agriculture Organization, 133,
Transgenics, 284 263
768
Subject Index
Urbanization; global warming, Warning coloration, 415

296; wildlife, 659, 666 Wasps, 344; as pollinators, 498
Urushiol, 493 Waste management, 667, 671;
tundra biomes, 657
Vacant niche theory, 645 Wastewater drainage, 225
Van Dyne, George, 192 Water; circulation in lakes, 367;
Variability in populations, 520 contamination, 501; in soil
Varves, 372 ecosystems, 597; wetlands, 672
Vascular tissues, 145, 241 Water column, 367, 393
Vaux’s swifts, 109 Water pollination, 499
Vavilov, Nikolai I., 279 Water table, 341
Vegetation corridors, 378 Watersheds, 440
Venoms, 486 Way, M. J., 115
Verhulst, Pierre-François, 577 Weathering and soil generation,
Vertical migrations, 409 594
Vervet monkeys, 98 Weeds, 356; allelopathy, 15
Vesicular-arbuscular mycorrhizae, Wetlands, 391, 672, 676
102, 426 Whales, overhunting of, 199
Vestiges of the Natural History of White, Gilbert, 98, 255
Creation (Chambers), 237 White pine blister rust, 41
Vestigial organs, 228 Whittaker, R. H., 57
Vicariance, 38 Whooping cranes, 203
Vinblastine, 404 Wildebeests, 388
Vincristine, 404 Wildlife. See Endangered species;
Vipers, 487; Gaboon, 488 Wildlife management; Zoos
Viruses as biopesticides, 65 Wildlife management, 143, 380,
Visual displays, 167, 359 512, 534, 677, 680; grasslands,
Vocalizations; birds, 97; and 301; urban and suburban, 665
ethology, 220 Wildlife preservation, 374
Volcanic activity and extinctions, Wildlife refuges, 203, 609, 678. See
247 also Nature preserves; Parks;
Volcanoes, 364 Zoos
Voles, red tree, 109 Wildlife, urban and suburban,
Volterra, V., 115 659, 666
Vomeronasal organs, 478 Wilson, Edward O., 38, 343, 578
Vries, Hugo de, 239 Wind erosion, 212
Vultures, 158 Wind pollination, 498
Winter, global warming, 293
Wallace, Alfred Russel, 38, 238 Wolves, 300; hierarchies, 331;
Warblers, competition among, 115 social organization, 389
Warm-blooded animals, 10 Worker ants, 343
769
Subject Index
World Conservation Union, 196, Zebrafish, 489

202 Zoological Philosophy (Lamarck),
World Health Organization, 503 179
Worldwatch Institute, 505 Zoology, ethology, 215
Worms as symbionts, 621 Zooplankton, 392, 483; eutrophic
Wright, Sewall, 522 environments, 223; migrations,
409
Xylem, 145 Zoos, 203, 439, 681, 685. See also
Nature preserves; Parks;
Yaks, introgression of, 84 Wildlife refuges
Yellowstone National Park, 316 Zosterophyllophyta (early land
Yews, 404; toxins, 492 plants), 241
Yucca, 156 Zygotic inviability, 360
Yucca Mountain, Nevada,
repository, 670
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What topics are covered in the ecology book?

What topics are covered in the ecology book?

What are some of the sections in Volume 1 of the book?

What are some of the sections in Volume 1 of the book?

Ecology Basics

The Editors of Salem Press

Mammalian social systems—Zoos

The Editors of Salem Press

Salem Press, Inc.

∞ The paper used in these volumes conforms to the American Na-

Library of Congress Cataloging-in-Publication Data

printed in the united states of america

Deep ecology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123

Ecology: definition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171

Food chains and webs . . . . . . . . . . . . . . . . . . . . . . . . . . . . 255

Gene flow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 274

Habitats and biomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313

Insect societies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 343

Lakes and limnology . . . . . . . . . . . . . . . . . . . . . . . . . . . . 364

Mammalian social systems . . . . . . . . . . . . . . . . . . . . . . . . . 385

Natural selection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 428

Ocean pollution and oil spills . . . . . . . . . . . . . . . . . . . . . . . . 444

Rain forests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 549

Savannas and deciduous tropical forests . . . . . . . . . . . . . . . . . 586

Urban and suburban wildlife . . . . . . . . . . . . . . . . . . . . . . . . 659

Waste management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 667

• Behavioral ecology: The study of how individual organisms interact

• Paleoecology: The study of past ecosystems and environments.

Steve K. Alexander Sneed B. Collard

Richard W. Arnseth J. A. Cooper

George K. Attwood Alan D. Copsey

David Landis Barnhill Mark S. Coyne

Erika L. Barthelmess Greg Cronin

Margaret F. Boorstein James F. Crow

Catherine M. Bristow John P. DiVincenzo

William R. Bromer Allan P. Drew

Jessica O. Ellison David Wason Hollar, Jr.

Danilo D. Fernando Robert Hordon

Robert Lovely Edward N. Nelson

Yiqi Luo Bryan Ness

Michael L. McKinney John G. New

Kristie Macrakis Edward B. Nuhfer

Paul Madden Oghenekome U. Onokpise

Nancy Farm Männikkö Robert W. Paul

Linda Mealey Rex D. Pieper

John S. Mecham Noreen D. Poor

Randall L. Milstein Robert Powell

Eli C. Minkoff Donald R. Prothero

Richard F. Modlin Carol S. Radford

Thomas C. Moon P. S. Ramsey

Randy Moore C. Mervyn Rasmussen

Ronald J. Raven Sanford S. Singer

Acid deposition, 1 Deforestation, 131

Geochemical cycles, 288 Human population growth, 333

Mammalian social systems, 385 Population analysis, 507

Trophic levels and ecological Wildlife management, 677

Types of ecology: Aquatic and marine ecology; Ecotoxicology

Activists blame coal-burning

In major cities, exhaust from automobiles combined with power plant

Effects on Aquatic Ecosystems