Contents
Foreword vii
Gary S. Hartshorn
Foreword ix
Luis Diego Gomez P.
A Manera de Introduccion xi
Luis Diego Gomez P.
Contributors xix
1. Introduction 3
Nalini M. Nadkarni & Nathaniel T. Wheelwright
1.1 Historical Overview 3
1.2 The Boundaries 5
Monteverde Beginnings 10
Wilford "Wolf" Guindon
Scope of Past Work 11
Nalini M. Nadkarni
Literature Cited 13
2. The Physical Environment 15
Kenneth L Clark, Robert 0, Lawton, & Paul R. Butler
2.1 Climate and Weather of Monteverde 15
2.2 Geology and Geologic History of Monteverde 20
2.3 Modern Geography of the Cordillera de Tilaran 22
2.4 Paleoecology of the Cordillera de Tilaran 26
2.5 Soils of Monteverde 27
2.6 Hydrology of Monteverde 29
Microclimate Variability 33
Kenneth L Clark & Nalini M. Nadkarni
Epiphytic Histosols 34
Kenneth L Clark & Nalini M. Nadkarni
Literature Cited 34
3. Plants and Vegetation
William A. Haber
39
3.1 Distribution and Diversity
3.2 Seasonality 64
3.3 Population Biology 66
3.4 Biogeography 67
3.5 Conservation 69
Trees on Trees 70
Francis E. Putz
Why Strangler Figs Don't Strangle Strangler Figs 71
Jonathan H. Titus
Mucuna urens, a Tropical Liana 72
Gary W. Diller O'Dell
Epiphytes 72
Stephan W. Ingram
Bromeliads 73
Harry E. Luther
Orchids 74
John T. Atwood
Evolution in Cloud Forest Psychotria
Section Notopleura via Growth Form
Diversification 75
Molly Nepokroeff& Kenneth J. Sytsma
Bryophytes 78
S. Robert Gradstein
Plants Growing on Living Leaves 80
Maria Morales
Mistletoes and Where They Grow 81
Sarah Sargent
An Exceptional Mistletoe, Gaiadendron
punctatum 82
Sarah Sargent
Gut Fungi of Invertebrates 83
Robert W. Lichtwardt
Breeding Systems of Monteverde Inga 85
Suzanne Koptur
Sex Ratios and the Distribution of
Male and Female Trees 87
Nathaniel T. Wheelwright
Clonal Reproduction through Plant
Fragments in Poikilacanthus
macranthus 88
Stephen P. Bush
Ferns 89
Seth Bigelow & Peter Kukle
Description of Permanent Forest Plots
in Monteverde 90
William A. Haber
Literature Cited 90
4. Insects and Spiders 95
Edited by Paul Hanson
4.1 Aquatic Insects 96
4.2 Orthoptera: Katydids, Crickets, and
Grasshoppers 98
4.3 Hemiptera: Heteroptera and
Homoptera 99
4.4 Coleoptera: Beetles 106
4.5 Lepidoptera: Moths and
Butterflies 118
4.6 Diptera: Flies 122
4.7 Hymenoptera: Sawflies, Wasps, Ants,
and Bees 124
4.8 Arachnids: Spiders, Scorpions, and
Mites 139
Literature Cited 141
5. Amphibians and Reptiles 149
J. Alan Pounds
5.1 The Area and Its Fauna 151
5.2 Distribution and Diversity 153
5.3 Geography and History 155
5.4 Population Ecology 157
5.5 Reproductive Ecology 163
5.6 Habitat Use 166
5.7 Conservation 169
5.8 Summary 170
The Discovery of the Golden Toad 171
Jay M. Savage
Monteverde Salamanders, Golden Toads,
and the Emergence of the Global Amphibian
Crisis 172
j. Alan Pounds
How Do Meadow Treefrogs Decide Where
to Lay Eggs? 173
Martha L Crump
The Golden Toad Laboratory for
Conservation 174
J. Alan Pounds & Marlene Brenes
Literature Cited 174
6. Birds 179
Bruce £. Young & David B. McDonald
6.1 Distribution, Species Richness, and
Diversity 182
6.2 Behavior 184
6.3 Reproductive Biology 190
6.4 Responses to Seasonality 194
6.5 Population Biology 195
6.6 Community Ecology 199
6.7 Comparative Ecology 201
6.8 Conservation Issues 202
6.9 Conclusions 203
Cooperation Between Male Long-Tailed
Manakins 204
David B. McDonald
Why Join Mixed-Species Flocks? A Frugivore's
Perspective 205
Lisa K. Valburg
The Cost of Social Foraging in Mixed-Species
Bird Flocks 206
Jennifer Shopland
Choosiness and Productivity in Wrens of
Forests, Fragments, and Farms 208
Kathy Winnett-Murray
Do Fruit-Eating Birds Actively Select or
Avoid Insect-Infested Fruits? 210
Lisa K. Valburg
XIV Contents
Brown Jays: Complex Sociality in a Colonizing
Species 212
Dean A. Williams & Marcy F. Lawton
Ecology and Sexual Dimorphism of Emerald
Toucanets 214
Cecilia M. Riley & Kimberly G. Smith
The Roles of Long-Tailed Manakin
Vocalizations in Cooperation and
Courtship 215
Jill M. Trainer
Do Subordinate Species Have an Advantage?
Testing the Pointer Hypothesis with Tropical
Hummingbirds 216
Harry M. Tiebout III
Literature Cited 218
7. Mammals 223
Robert M. Timm & Richard K. LaVal
7.1 Methods 223
7.2 Distribution, Species Richness, and
Diversity 224
7.3 Research on Mammals in
Monteverde 225
7.4 Migration 229
7.5 Changes in Altitudinal Limits of Life
Zones as Suggested by Bats 231
7.6 Historical Use and Change in
Abundance of Mammals 232
7.7 Conservation of Monteverde
Mammals 234
Observations on Monteverde's Mammals 235
Robert M. Timm & Richard K. LaVal
Singing Mice 236
Catherine A. Langtimm
Reproduction and Dynamics of Deer Mice 238
Stephen Anderson
Arboreal Mammals 239
Catherine A. Langtimm
Mice, Birds, and Pollination of Blakea
chlorantha 241
Catherine A. Langtimm & Robert Unnasch
Baird's Tapir 242
Robert 0. Lawton
Literature Cited 243
8. Plant-Animal Interactions 245
K. Greg Murray, Sharon Kinsman,
& Judith L Bronstein
8.1 Plant-Pollinator Interactions 246
8.2 Plant-Frugivore Interactions 256
Hummingbird Pollination of Epiphytic
Ericaceae in the Cloud Forest Canopy 267
William H. Busby
Scarab Beetles, Elephant Ear (Xanthosoma
robustum), and Their Associates 268
Lloyd Goldwasser
Fig Pollination and Seed-Dispersal
Mutualisms 271
Judith L Bronstein
The Reproductive Biology of Blakea and
Topobea (Melastomataceae) 273
Cecile Lumer
Interactions among Inga, Herbivores, Ants, and
Insect Visitors to Foliar Nectaries 277
Suzanne Koptur
A Fly Larva Directly Alters Floral Sex in
Centropogon solanifolius 278
Martha E, Weiss
Deceit Pollination in Begonia 279
Jon Agren & Douglas W. Schemske
A Hypothesis about the Timing of Flowering
and Fruiting in Competing Tropical Trees 281
Nathaniel T. Wheelwright
Fruiting Phenologies of Pioneer Plants:
Constraints Imposed by Flowering Phenology,
Disturbance Regime, and Disperser Migration
Patterns 283
K. Greg Murray
What Happens to Seeds of Vertebrate-
Dispersed Trees after Dispersal? 286
Dan Wenny
Specialized Seed Dispersal: Mistletoes and
Fruit-Eating Birds 288
Sarah Sargent
Seed Dispersal and Seedling Recruitment in a
Lauraceous Tree Species 289
J. P. Gibson
Myrmecophytes 291
John T. Longino
The Importance of Different Bird Species as
Seed Dispersers 294
K. Greg Murray
Literature Cited 295
9. Ecosystem Ecology and Forest Dynamics 303
Nalini M. Nadkarni, Robert 0. Lawton,
Kenneth L Clark, TeriJ. Matelson,
& Doug Schaefer
9.1 Forest Structure, Composition, and
Dynamics 304
9.2 Forest Biogeochemistry and Nutrient
Cycling 320
9.3 Future Research Directions 335
Microbial Biomass and Activity in Canopy
Organic Matter and the Forest Floor 336
Eric Vance & Nalini M. Nadkarni
XV Contents
Invertebrates in Canopy and Ground Organic
Matter 336
Nalini M. Nadkarni &John T. Longino
Vesicular-Arbuscular Mycorrhizae of Epiphytic
and Terrestrial Piperaceae 338
Blase Maffio, Nalini M. Nadkarni,
& David P. Janos
Factors Affecting the Initiation and Growth
of Aboveground Adventitious Roots in a
Tropical Cloud Forest Tree: An Experimental
Approach 339
Nalini M. Nadkarni
Fine Litter Dynamics within the Tree Canopy
of a Tropical Cloud Forest 341
Nalini M. Nadkarni & Teri J. Matelson
Longevity of Fallen Epiphytes 344
Teri j. Matelson, Nalini M. Nadkarni,
&John T. Longino
Literature Cited 345
10. Conservation in the Monteverde Zone:
Contributions of Conservation
Organizations 351
Leslie], Burlingame
10.1 The Context 352
10.2 The Quakers and Bosqueterno,
S.A. 356
10.3 The Monteverde Cloud Forest
Preserve 357
10.4 The Monteverde Conservation League
and Bosque Eterno de los Nifios 361
10.5 Santa Elena High School Cloud Forest
Reserve 367
10.6 The Monteverde Institute 369
10.7 Environmental Education in Public
and Private Schools 370
10.8 Conclusion: Lessons from Monteverde
and Topics for Future Research 372
Key to Abbreviations Used in this
Chapter 375
The Changing Face of Tourism 376
Sam Crosby
Pros and Cons of Ecotourism 376
Francisco Chamberlain
Debt-for-Nature Swaps 377
Leslie]. Burlingame
The Community Process of Environmental
Education 377
Guillermo Vargas (Translated by Leslie J.
Burlingame)
Monteverde 2020 378
Leslie]. Burlingame
El Buen Amigo 379
Eugenio Vargas
La Bella Farm 380
Leslie]. Burlingame
San Gerardo: An Experiment in Sustainable
Development 380
John Boll
Conservation Easements in Monteverde: The
Enlace Verde Project 381
Nathaniel Scrimshaw, Wendy Gibbons,
& Leslie]. Burlingame
The Monteverde Butterfly Garden 382
Jim Wolfe
Comite de Artesanias Santa Elena-
Monteverde 383
Leslie]. Burlingame
La Campesinita 384
Leslie]. Burlingame
Human Voices around the Forest 385
Eugenio Vargas
Literature Cited 385
11. Agriculture in Monteverde: Moving Toward
Sustainability 389
Katherine Griffith, Daniel Peck, & Joseph Stuckey
11.1 The Monteverde Dairy Industry 392
11.2 Lowland Dual-Purpose and Beef
Production 398
11.3 Coffee Production 400
11.4 Other Crops 403
11.5 Livestock and Fowl 405
11.6 Future Directions and Conclusions 405
Prehistoric Cultures and Inhabitants 408
Robert M. Timm
The Agroecology of Prosapia: Spittlebugs,
Froghoppers, and Pasture Pests 409
Daniel Peck
Brassicas as Biological Plows 411
Jim Wolfe
Pasture Burning 411
Katherine Griffith
Monteverde's Kitchen Gardens 412
Katy VanDusen
Pesticide Use in the Santa Elena Area 414
Katherine Griffith
Frijol Tapado or "Covered Beans" 414
Katherine Griffith &Jim Wolfe
The Pig Farm: "Sustainable Agriculture" or "an
Abomination"? 415
Katherine Griffith
Literature Cited 416
xvi Contents
12. Conservation Biology 419
Nathaniel T. Wheelwright
12.1 Richness, Endemism, and
Biodiversity 420
12.2 An Overview of Key Concepts in
Conservation Biology 420
12.3 The Status of Biodiversity in
Monteverde: What Is at
Stake? 423
12.4 Conservation Problems in
Monteverde 424
12.5 Can Conservation Biology Offer
Solutions to Monteverde's
Conservation Problems? 428
12.6 Conclusions 431
Enduring Reasons to Preserve Species 432
Nathaniel T. Wheelwright
How Have Humans Affected Bird
Populations? 433
Bruce E. Young
The Influence of Fruit-Eating Bats on the
Dynamics and Composition of Neotropical
Premontane Cloud Forests 434
Eric Dinerstein
The Importance of Pacific Slope Forest for
Maintaining Regional Biodiversity 435
Carlos F. Guindon
La Ventana in Monteverde: A Migration
Corridor for Insects 438
Michelle Williamson & Chris Darling
Altitudinal Migrations and Habitat Linkages:
Using the Resplendent Quetzal as an Indicator
for Evaluating the Design of the Monteverde
Reserve Complex 439
George V. N. Powell, Robin D. Bjork,
Sergio Barrios, & Vicente Espinoza
Patterns in the Regeneration of Lauraceous
Trees in Abandoned Pastures 442
Martha Groom
The Impact of the Monteverde Lecheria on the
Quebrada Guacimal 446
Douglas E. Gill
House Wrens in Monteverde: A Population
Sink? 448
Bruce E. Young
Windbreaks as Corridors for Birds 448
Karen Nielsen & Debra DeRosier
Windbreaks as Habitats for Trees 450
Celia A. Harvey
The Search for Medicines from the Plants of
Monteverde 452
William N. Setzer
Literature Cited 453
Appendixes 457
Appendix 1. Vascular Plants of Monteverde 457
William A. Haber
Appendix 2. Number of Species with Different
Plant Growth Forms 519
William A. Haber
Appendix 3. Orchids of Monteverde 523
John 7". Atwood
Appendix 4. Bromeliads of Monteverde 529
Harry £ Luther
Appendix 5. Host Plants of Selected Cercopoidea
from Monteverde 531
Daniel C Peck
Appendix 6. Preliminary List of Psylloidea
Known from Costa Rican Cloud Forests
(1200-2000 m) 533
David Hollis
Appendix 7. Eusocial Wasps (Hymenoptera: Vespidae
Polistinae and Spheddae) Collected from
Monteverde 535
Sean O'Donnell
Appendix 8. Amphibians and Reptiles of
Monteverde 537
J. Alan Pounds & M. P. Fogden
Appendix 9. Birds of the Monteverde Area 541
M. P. Fogden
Appendix 10. Mammals of Monteverde 553
Robert M. Timm & Richard K. LaVal
References 559
Index 561
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