Table Of ContentFOOD WEBS
Integration 0& Patterna & Dynamica
FOOD WEBS
Integration o~ Pattern& (1 Dynamic&
Edited by
Gar)' A. PoLi&
Department of Biology, Vanderbilt University, Nashville, Tennessee
Kirk O. Winemiller
Department of Wildlife and Fisheries, Texas A & M University, College Texas
SPRINGER-SCIENCE+BUSINESS MEDIA, B.V.
Copyright © 1996 by Springer Science+Business Media Dordrecht
Originally published by Chapman & Hall in 1996
Softcover reprint of the hardcover 1st edition 1996
All rights reserved. No part ofthis book covered by the copyright hereon may be reproduced or used in any form or by any means--graphic,
electronic, or mechanical, inciuding photocopying, recording, taping, or information storage and retrieval systems--without the written
pennission ofthe publisher.
2 3 4 5 6 7 8 9 XXX 01 00 99 98 97 96
Library of Congress Cataloging-in-Publication Data
Food webs: integration ofpatterns and dynamics / [edited by] G. A. Polis and K. O. Winemiller.
p. cm.
Includes bibliographical references and index.
ISBN 978-1-4615-7009-7 ISBN 978-1-4615-7007-3 (eBook)
DOI 10.1007/978-1-4615-7007-3
I. Food chains (Ecology) I. Polis, Gary A., 1946- 11. Winemiller, K. O. (Kirk 0.), 1956--
QH54l.F66 1995 94-43155
574.5"3--dc2 CIP
Table of Contents
Preface
Robert T. Paine ix
List of Contributors xi
Introduction Food Webs: What Do They Tell Us About the World?
Kirk O. Winerniller and Gary A. Polis
SECTION I Detritus and Nutrients
1. Detritus and Nutrients in Food Webs 25
Michael f. Vanni and Peter C. DeRuiter
2. Food Webs and Nutrient Cycling in Soils: Interactions and Positive Feedbacks 30
fanne Bengtsson, Heikki Setälä, and D. W. Zheng
3. Energetics of Detritivory and Microbivory in Soil in Theory and Practice 39
David C. Colernan
4. Integrating the Microbial Loop and the Classic Food Chain into a Realistic 51
Planktonic Food Web
Karen G. Porter
5. Trophic Structure and Carbon Flow Dynamics in the Pelagic Community of a 60
Large Lake
Ursula Gaedke, Dietrnar Straile, and Claudia Pahl-Wostl
6. Biogeochemistry and Trophic Ecology: A New Food Web Diagram 72
Robert W. Sterner, farnes f. Elser, Thomas H. Chrzanowski, fohn H. Scharnpei,
and Nicolas B. George
7. Nutrient Transport and Recycling by Consumers in Lake Food Webs: Implications 81
for Algal Communities
Michael f. Vanni
8. Food Web Structure and Littoral Zone Coupling to Pelagic Trophic Cascades 96
Daniel E. Schindler, Stephen R. Carpenter, Kathryn L. Cottingharn, Xi He,
farnes R. Hodgson, farnes F. Kitchell, and Patricia A. Soranno
SECTION 11 Interaction of Productivity and Consumption
9. Interaction of Productivity and Consumption 109
Donald L. DeAngelis, Lennart Persson, and Arny D. Rosernond
10. Dynamics and Interactions in Food Webs with Adaptive Foragers 113
Peter A. Abrarns
v
vi / Table 0/ Contents
11. Nonlinear Food Web Models and Their Responses to Increased Basal Productivity 122
Roger Arditi and Jerzy Michalski
12. The Relative Importance of Resource Limitation and Predator Limitation in 134
Food Chains
Craig W. Osenberg and Gary G. Mittelbach
13. Indirect Effects of Herbivores Modify Predicted Effects of Resources and 149
Consumption on Plant Biomass
Amy D. Rosemond
14. Food Web Dynamics on Some Small Subtropical Islands: Effects of Top and 160
Intermediate Predators
David A. Spiller and Thomas W. Schoener
15. Top Down from Underground? The Underappreciated Influence of Subterranean 170
Food Webs on Aboveground Ecology
Donald R. Strong, John L. Maron, and Peter G. Connors
SECTION III Causes and Effects
16. Causes and Effects in Food Webs: Do Generalities Exist? 179
Janne Bengtsson and Neo Martinez
17. Assessing the Relative Importance of Trophic Links in Food Webs 185
David G. Raffaelli and Stephen J. Hall
18. Food Webs and Perturbation Experiments: Theory and Practice 192
Peter Yodzis
19. Energetics and Stability in Belowground Food Webs 201
Peter C. DeRuiter, Anje-Margriet Neutel, and John C. Moore
20. What Equilibrium Behavior of Lotka-Volterra Models Does Not Tell Us About 211
Food Webs
Alan Hastings
21. Effects of Food Chain Length and Omnivory on Population Dynamics in 218
Experimental Food Webs
Peter J. Morin and Sharon P. Lawler
22. Structure and Dynarnics of Arctic-Subarctic Grazing Webs in Relation to 231
Primary Productivity
Lauri Oksanen, Tarja Oksanen, Per Ekerholm, Jon Moen, Peter Lundberg,
Michael Schneider, and Maano Aunapuu
23. Food Webs: From the Lindeman Paradigm to a Taxonomic General Theory 243
of Ecology
Steven H. Cousins
SECTION IV Temporal and Spatial Sc ale
24. Temporal and Spatial Aspects of Food Web Structure and Dynamics 255
Robert D. Holt
25. Control of Interaction Strength in Marine Benthic Communities 258
Bruce A. Menge, Bryon Daley, and P. A. Wheeler
26. Allochthonous Input Across Habitats, Subsidized Consumers, and Apparent Trophic 275
Cascades: Examples from the Ocean-Land Interface
Gary A. Polis and Stephen D. Hurd
27. Disturbance and Food Chain Length in Rivers 286
Mary E. Power, Michael S. Parker, and J. Timothy Wootton
28. Factors Driving Temporal and Spatial Variation in Aquatic Floodplain Food Webs 298
Kirk O. Winemiller
29. Food Webs in Space: An Island Biogeographic Perspective 313
Robert Holt
Table of Contents / vii
SECTION V Food Webs and Applied Problems
30. Food Web Dynamics and Applied Problems 327
Larry B. Crowder, Douglas P. Reagan, and Diana W. Freckman
31. Structure and Impact of Natural Enemy Guilds in Biological Control of Insect Pests 337
Les E. Ehler
32. Quantifying the Impact of Insecticides on Food Web Structure of Rice Arthropod 343
Populations in a Philippine Farmer's Irrigated Field: A Case Study
Kenneth G. Schoenly, Joel E ..C ohen, K. L. Heong, Gertrudo S. Arida,
Alberto T. Barrion, and James A. Litsinger
33. The Impact of Industrial Fisheries on the Trophic Structure of Marine Ecosystems 352
Timothy R. Parsons
34. Trophic Flow Networks as Indicators of Ecosystem Stress 358
Robert E. Ulanowicz
SECTION VI Syntheses
35. The Role of Indirect Effects in Food Webs 371
Peter Abrams, Bruce A. Menge, Gary G. Mittelbach, David Spiller, and
Peter Yodzis
36. Productivity and Consumer Regulation-Concepts, Patterns, and Mechanisms 396
Lennart Persson, Janne Bengtsson, Bruce A. Menge, and Mary E. Power
37. Time, Space, and Life History: Inftuences on Food Webs 435
Gary A. Polis, Robert D. Holt, Bruce A. Menge, and Kirk O. Winemiller
Index 461
Preface
Food webs remain the ecologically flexible what units (species, trophospecies, kjoules,
scaffolding around which communities are nutrients) were the most suitable, what the
assembled and structured. Furthermore, webs appropriate spatial and temporal scales were,
suggest the pathways by which materials are and whether a static approach could be mean
transferred and along which energy flows. ingfully applied to a dynamic natural world.
As such, they are fundamental and in many One milestone, the 1982 conference "Cur
senses inescapable in any attempt to describe rent Trends in Food Web Theory, hosted by
how natural assemblages are organized or scientists at the Oak Ridge National Lab,
complexes of species interact. Although the reflected contemporary beliefs and interests.
roots go historically much deeper, Darwin's The present volume is a worthy successor
"tangled bank" metaphor is a convenient, and suggests both a decade of conceptual
qualitative start. More modem landmarks evolution and a relatively painless refocusing
would be Charles Elton's 1927 book-essen of the issues. Advances have come primarily
tially marking the birth of community ecol from recognizing the underworld, the arena
ogy-and Raymond Lindeman 's 1942 paper of nutrient regeneration and decomposers.
on trophic dynamics signaling the appearance The basic problems remain daunting, how
of energy units as ecological currency. If one ever: how to recognize links; the necessity
accepts this simplistic overview, many of the for incorporation of nontrophic relationships,
intellectual trends and academic alliances especially interference competition; and how
characterizing North American and European to confront the spatial and temporal variabil
ecology fall into distinctive but not exclusive ity of nature, which involves a cast of thou
cults. sands of thousands of species, most of whose
An impressive proportion of what I believe roles remain unknown and unappreciated.
to have been ecology's recent bandwagons There clearly are numerous hurdles to be
relate to food webs. The stability-diversity surmounted, but perhaps the major challenge
interplay was the first, and energetics and the is to avoid succumbing to the almost irresist
International Biological Program were close ible temptation of overdefinition of a complex
on its heels. These issues dominated two de phenomenon. As an example, no one doubts
cades beginning about 1955. There were that ecological assemblages change through
then, starting about 1970, 15-20 years of time. Frederick Clements developed succes
food web numerology during which intense sion as ecology's first paradigm, and then
effort was made to assemble representative proceeded to bury this nascent area in a load
and respectable webs and to analyze them of unnecessary and counterproductive jar
for emergent patterns. Patterns were found, gon. One can draw paralleis with food webs,
creative analyses published, and the whole likewise a derived description of complex
approach challenged, principally on the issue interactions. Excessive coining of new terms
of what links were significant or spurious, will similarly render the current effort to un-
ix
x / Preface
derstand the compIexities of trophic relation a litany oftrophic ecology sins to be avoided.
ships relatively uninterpretable a few years I believe this symposium has succeeded ad
hence, just like most of Clements' and related mirably on most counts: manipulation is seen
descriptive literature is uninterpretable now, as the handmaiden of description; decom
even with a dictionary of terms. posers and nutrients have begun to assurne a
If resisting the temptation to coin provoca central position; the vexing problems associ
tive yet descriptive vocabulary is the first ated with temporal and spatial scaling are
commandment of food web research ("thou openly acknowledged; the dynamic conse
shalt not commit jargon"), what might be the quences of interspecific interactions are in
second? Although precise wording escapes creasingly apparent (meaning webs are per
me, I believe it should develop the admoni mitted, even expected, to change from
tion that the structure of nature guide mathe within, rather being forced by externalities):
matical development and not vice versa. "web metrics," perhaps like their ill-fated
Thus, most naturalists know omnivory to be predecessor "niche metrics," are pIeasantly
commonplace. Why develop models chal inconspicuous.
Ienging or obscuring that fact? The third and What I have written represents ablend of
later commandments might explore the im long-held personal prejudices and a selective
portance of temporal variability and spatial examination of chapters, titles, and summar
scale as inftuences in observable trophic pat ies in the book. Food web research has moved
terns, and the primary necessity of incorpo a great intellectual distance since 1982, let
rating dynamics. Without acknowledging alone 1942 or 1927. One cannot describe
this central feature of all known assemblages communities or ecosystems without em
(that is, that the component species interact), ploying interunit connections. Food webs,
there is littIe hope for the sorts of local pre liberally interpretated, provide this frame
dictive capability useful to conservationists work and are thus unescapable. Creative
and ecosystem managers. thought on webs, richly evident in the last
One might also hope for a statement on the decade, supports my optimism that the holy
importance of species (let alone individuals): trinity of field observation, controlled experi
"thou shalt honor species individuality." mentation, and mathematical exploration can
Such aggregations as guilds, trophospecies, discover some common unity, and in the pro
even trophic levels themselves, perhaps all cess provide important guidelines on how to
necessary to minimize nature' s complexity, save and even resurrect our increasingly bat
also serve to isolate web members from the tered natural world. Such a capability is the
details of behavioral biology and the deep litmus test of ecological value; food webs are
insights of natural selection. Finally, per central to such endeavors.
haps, should be an admonition not to exclude
the taxonomically awkward soil and meiofau ROBERT T. PAINE
nal organisms, picoplankton, fungi, and Department of Zoology
other decomposers, and their little known in University of Washington
teractions. Seattle, Washington
Such a list is hardly exhaustive but it covers 98195
List of Contributors
Peter A. Abrams Thomas H. Chrzanowski
Department of Ecology, Evolution and Department of Biology
Behavior Box 19498
University of Minnesota University of Texas at Arlington
1987 Upper Buford Circle Arlington, TX 76019 USA
St. Paul, MN 55108 USA
Joel E. Cohen
Laboratory of Populations
Roger Arditi
Box 20
Institute of Zoology and Animal Ecology
RockefeIler University
University of Lausanne
1230 York Avenue
CH-1015 Lausanne, Switzerland
New York, NY 10021 USA
Gertrudo S. Arida David C. Coleman
Entomology and Plant Pathology Division Institute of Ecology
International Rice Research Institute University of Georgia
Los Banos, Philippines 711 Biological Sciences Building
Athens, GA 30602 USA
Maano Aunapuu
Peter G. Connors
University of Tartu
University of Califomia, Davis
Vanemuise 46
Bodega Bay Marine Laboratory
EE-2400 Tartu, Estonia
Box 247
Bodega Bay, CA 94923 USA
Alberto T. Barrion
Entomology and Plant Pathology Division Kathryn L. Cottingham
International Rice Research Institute Center for Limnology
Los Banos, Philippines 680 N. Park Street
Madison, WI 53706 USA
Janne Bengtsson
Steven H. Cousins
Section for Soil Ecology
International Ecotechnology Research
Department of Ecology and
Centre
Environmental Research
Cranfield University
Swedish University of Agricultural
Cranfield Bedfordshire MK43 OAL UK
Sciences
Box 7072, S-750 07 Uppsala, Sweden
Larry B. Crowder
Duke University School of the
Stephen R. Carpenter, Environment
Center for Limnology Marine Laboratory
680 N. Park Street 135 Duke Marine Lab Road
Madison, WI 53706 USA Beaufort, NC 28517-9721 USA
xi
