THE FOOD WEB OF A TROPICAL RAIN FOREST
Edited by Douglas P. Reagan and Robert B. Waide
The University of Chicago Press / Chicago and London
1996
CONTENTS
Preface
1 The Rain Forest Setting
Robert B. Waide and Douglas P. Reagan
A Tropical Forest Community 3
The Biogeographical Context of the Food Web at El Verde 4
The Ecological Context of the Food Web at El Verde 6
The El Verde Food Web 11
The Study of Feeding Relationships in Animal Communities 13
2 Plants: The Food Base
William T. Lawrence, Jr.
The Energetic Basis of the Food Web 18
Diversity of the Flora 20
Disturbance Effects 21
Plant Biomass and Productivity 21
Leaf, Flower, and Fruit Phenology 30
Controls on Primary Consumption 34
Summary 39
Appendix 2: Plants at El Verde 41
3 Microorganisms
D. Jean Lodge
Taxonomic Surveys 55
Density and Biomass 55
Effects of Disturbance on Fungi 60
Microorganisms in the Food Web 62
Linkages Involving Microorganisms in the Food Web 68
Microbial Processes: Rates and Flows 74
Summary 78
Acknowledgments 80
Appendix 3.A: Partial Checklist of Fungi 81
Appendix 3.B: Presence of Infections 103
Appendix 3.C: Mycorrhizal Classification and Methods 105
4 Termites
Elizabeth A. McMahan
Taxonomic Affinities of El Verde Termites 110
Population Density and Biomass 121
Diet 124
Consumption Rates and Energy Flows (Population Energetics) 129
Summary 134
5 Litter Invertebrates
William J. Pfeiffer
Dynamics of Litter Moisture and Standing Crop 138
Disturbance and the Litter Invertebrate Community 140
Litter and Soil Foraging Guilds 141
Vertebrate Predation on Litter Arthropods 175
Consumption Rates 175
Summary 177
Appendix 5: Study Site and Sampling Methods 180
6 Arboreal Invertebrates
Rosser W. Garrison and Michael R. Willig
History 186 187
Taxonomic Status of Invertebrates at El Verde 187
Diversity 190
Density 192
Post-Hurricane Effects 194
Age Structure 195
Seasonality 196
Feeding Guilds 197
Foraging Activity 207
Vertical Stratification 211
Energy Flow and Nutrient Cycling 213
Summary 214
Acknowledgments 215
Appendix 6: Foraging Status of Invertebrates in the El Verde Rain
Forest 217
7 Arboreal Arachnids
William J. Pfeiffer
Community Structure 248
Arachnids in the Invertebrate Community 252
Spider Populations of the Arboreal Strata 253
Other Understory Arachnids 265
Summary 267
Appendix 7: Methods 270
8 Amphibians
Margaret M. Stewart and Lawrence L. Woolbright
Methods 276
Natural History 278
Frogs as Predators 287
Frogs as Prey 302
Energy and Nutrient Cycling 309
Summary 314
Acknowledgments 315
Appendix 8.A: List of Prey Taxa 317
Appendix 8.B: Methods 320
9 Anoline Lizards
Douglas P. Reagan
Ecological Distribution 323
Populations 329
Food Habits 334
Predators 343
Summary 343
Acknowledgments 345
10 Nonanoline Reptiles
Richard Thomas and Ava Gaa Kessler
Microhabitats 350
Predation 354
Food Hahits 355
Populations and Biomass 360
Summary 361
Acknowledgments 362
11 Birds
Robert B. Waide
Distribution and Affinities of the El Verde Avifauna 364
Annual and Seasonal Changes in Avian Populations 370
Community Structure 376
Diet 381
Foraging Distribution 387
Birds in the El Verde Food Web 390
Summary 394
Acknowledgments 396
Appendix 11: Methods 397
12 Mammals
Michael R. Willig and Michael R. Gannon
Zoogeographic Considerations 400
Bats 403
Terrestrial Consumers 419
Response to Disturbance 422
Trophic Comparisons 425
Discussion 427
Summary 429
Acknowledgments 430
13 The Stream Community
Alan P. Covich and William H. McDowell
Physical Controls on Stream Food Webs 435
Biotic Controls on Stream Food Webs 442
Feeding Groups 450
Connections with Terrestrial Food Webs 454
Patterns of Feeding Relationships 456
Summary 459
14 The Community Food Web: Major Properties and Patterns of Organization
Douglas P. Reagan, Gerardo R. Camilo, and Robert B. Waide
Characteristics of the El Verde Animal Community 462
Attributes of the Food Web 470
Conclusions 486
Summary 487
Appendix 14.A: The Food Web Matrix 489
Appendix 14.B: Mean Annual Populaton Densities and Biomass for Different
Taxa 508
Glossary
Contributors
Bibliography
Index
FROM THE BACK COVER
BUILDING ON more than a decade of field research, this volume presents a
comprehensive description and analysis of the tropical rain forest at El
Verde, Puerto Rico. The contributors weave the strands of information about
the energy flow within the forest into the most complete and detailed
description of a complex food web ever attempted. The results shed new
light not only on this community, but also on the principles guiding the
organization of all food webs.
After an introduction describing the background and environmental setting
of the study, twelve chapters address in detail the specific organisms that
inhabit El Verde: plants, microorganisms, invertebrates, frogs, reptiles,
birds, mammals, and various aquatic organisms in forest streams. Each
chapter summarizes information on the abundance, food habits, and
ecological roles of the organisms it covers, and places them within the
context of the wider community. Appendixes to these chapters list all
species discovered to date in the forest. The final chapter presents
conclusions regarding food web organization based on analyses of the
combined data, providing unique insights into the existence of food web
compartments, the role of reciprocal predation (food loops), and the degree
of omnivory in the community. The complete food web matrix used for the
analysis is included in an appendix, and an extensive bibliography contains
more than 1,500 references on the El Verde ecosystem and other relevant
literature.
This book is a valuable contribution to ecological knowledge, both for its
extended analysis of this Caribbean ecosystem and for its examination of
general principles of food web dynamics. Its systematic approach to
organizing the natural histories of the many species at El Verde reveals
basic patterns and processes common to all tropical rain forests, making
this book essential for anyone concerned with studying and protecting these
fragile ecosystems.
"This book represents a unique effort to study all the species in a
tropical rain forest, their natural histories, and the interconnections
among them. It is the most complete description of a food web yet produced."
-Stuart L. Pimm, author of Food Webs and The Balance of Nature?
"The Food Web of a Tropical Rain Forest is a unique and quite valuable
contribution to the field of ecology. It is the only book explicitly
organized to depict and then analyze the food web of an entire working
community, and should stand as a model for construction of realistic food
webs of other communities."
-Gary A. Polis, co-editor of Food Webs: Integration of Patterns and Dynamics
DOUGLAS P. REAGAN is a senior consultant at Woodward-Clyde Consultants in
Denver. ROBERT B. WAIDE is director of the Terrestrial Ecology Division at
the University of Puerto Rico.
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