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frankie front 8/20/03 7:44 PM Page i BIODIVERSITY CONSERVATION in COSTA RICA frankie front 8/20/03 7:44 PM Page ii frankie front 8/20/03 7:44 PM Page iii BIODIVERSITY CONSERVATION in COSTA RICA Learning the Lessons in a Seasonal Dry Forest Edited by Gordon W. Frankie, Alfonso Mata, and S. Bradleigh Vinson UNIVERSITY OF CALIFORNIA PRESS Berkeley Los Angeles London frankie front 8/20/03 7:44 PM Page iv University of California Press Berkeley and Los Angeles, California University of California Press, Ltd. London, England © 2004 by The Regents of the University of California Library of Congress Cataloging-in-Publication Data Biodiversity conservation in Costa Rica : learning the lessons in a seasonal dry forest / edited by Gordon W. Frankie, Alfonso Mata, and S. Bradleigh Vinson. p. cm. Includes bibliographical references. ISBN 0-520-22309-8 (cloth : alk. paper).—ISBN 0-520-24103-7 (pbk. : alk. paper). 1. Biological diversity conservation—Costa Rica. 2. Forest ecology— Costa Rica. I. Frankie, G. W. II. Mata, Alfonso. III. Vinson, S. Bradleigh, 1938– QH77.C8 B56 2004 333.95′16′097286—dc21 2003000593 Manufactured in the United States of America 13 12 11 10 09 08 07 06 05 04 10987654 321 The paper used in this publication meets the minimum requirements of ANSI/NISO Z39.48-1992 (R 1997) ∞ frankie front 8/20/03 7:44 PM Page v CONTENTS Preface / vii 4. IMPACT OF GLOBAL CHANGES ON THE REPRODUCTIVE BIOLOGY OF TREES IN 1. INTRODUCTION / 1 TROPICAL DRY FORESTS / 38 Alfonso Mata and Jaime Echeverría Kamaljit S. Bawa Part I. Biodiversity and 5. TROPICAL DRY-FOREST MAMMALS OF Ecological Studies PALO VERDE: ECOLOGY AND CONSERVATION IN A CHANGING LANDSCAPE / 48 Section A. Costa Rican Dry Forest Kathryn E. Stoner and Robert M. Timm 2. FLOWERING PHENOLOGY AND POLLINATION SYSTEMS DIVERSITY IN THE SEASONAL 6. THE CONSERVATION VALUES OF DRY FOREST / 17 BEES AND ANTS IN THE COSTA RICAN DRY FOREST / 67 Gordon W. Frankie, William A. Haber, S. Bradleigh Vinson, Kamaljit S. Bawa, S. Bradleigh Vinson, Sean T. O’Keefe, and Peter S. Ronchi, and Nelson Zamora Gordon W. Frankie 3. BREEDING STRUCTURE OF NEOTROPICAL 7. ECOLOGY OF DRY-FOREST WILDLAND DRY-FOREST TREE SPECIES IN INSECTS IN THE AREA DE FRAGMENTED LANDSCAPES / 30 CONSERVACIÓN GUANACASTE / 80 James L. Hamrick and Victoria J. Apsit Daniel H. Janzen frankie front 8/20/03 7:44 PM Page vi Section B. Biotic Relationships with Other Part II. Transferring Biodiversity Costa Rican Forests Knowledge into Action: The Record 8. DIVERSITY, MIGRATION, AND CONSERVATION OF BUTTERFLIES IN NORTHERN COSTA RICA / 99 17. BIODIVERSITY INVENTORIES IN COSTA RICA AND William A. Haber and Robert D. Stevenson THEIR APPLICATION TO CONSERVATION / 229 Paul Hanson 9. WATERSHED ECOLOGY AND CONSERVATION: HYDROLOGICAL RESOURCES IN THE 18. CONFLICT RESOLUTION: RECOGNIZING NORTHWEST OF COSTA RICA / 115 AND MANAGING DISCORD IN Alfonso Mata RESOURCE PROTECTION / 237 Gregory A. Giusti 10. WHERE THE DRY FOREST FEEDS THE SEA: THE GULF OF NICOYA ESTUARY / 126 19. CONSERVATION AND ENVIRONMENTAL José A. Vargas and Alfonso Mata EDUCATION IN RURAL NORTHWESTERN COSTA RICA: LEARNING THE LESSONS OF A NONGOVERNMENTAL ORGANIZATION / 247 11. MANGROVE FORESTS UNDER DRY SEASONAL CLIMATES IN COSTA RICA / 136 Gordon W. Frankie and Jorge A. Jiménez S. Bradleigh Vinson Section C. Biotic Relationships with Other 20. THE MEDIA AND BIODIVERSITY Geographical Areas CONSERVATION / 257 12. GEOGRAPHICAL DISTRIBUTION, ECOLOGY, Gilda Aburto AND CONSERVATION STATUS OF COSTA RICAN DRY-FOREST AVIFAUNA / 147 21. THREATS TO THE CONSERVATION OF TROPICAL DRY FOREST IN COSTA RICA / 266 Gilbert Barrantes and Julio E. Sánchez Mauricio Quesada and Kathryn E. Stoner 13. AN ULTRASONICALLY SILENT NIGHT: THE TROPICAL DRY FOREST WITHOUT BATS / 160 22. ENVIRONMENTAL LAW OF COSTA RICA: DEVELOPMENT AND ENFORCEMENT / 281 Richard K. LaVal Roxana Salazar 14. BIODIVERSITY AND CONSERVATION OF MESOAMERICAN DRY-FOREST 23. DISPUTE OVER THE PROTECTION OF THE HERPETOFAUNA / 177 ENVIRONMENT IN COSTA RICA / 289 Mahmood Sasa and Federico Bolaños Julio Alberto Bustos 15. PARQUE MARINO LAS BAULAS: CONSERVATION 24. THE POLICY CONTEXT FOR CONSERVATION IN LESSONS FROM A NEW NATIONAL PARK AND COSTA RICA: MODEL OR MUDDLE? / 299 FROM 45 YEARS OF CONSERVATION OF SEA TURTLES IN COSTA RICA / 194 Katrina Brandon James R. Spotila and Frank V. Paladino 25. CONCLUSION AND RECOMMENDATIONS / 311 16. PROSPECTS FOR CIRCA SITUM TREE Gordon W. Frankie, Alfonso Mata, and CONSERVATION IN MESOAMERICAN Katrina Brandon DRY-FOREST AGRO-ECOSYSTEMS / 210 David H. Boshier, James E. Gordon, List of Contributors / 325 and Adrian J. Barrance Index / 327 frankie front 8/20/03 7:44 PM Page vii PREFACE he idea of producing this book resulted lished in Costa Rica, including the dry-forest area, T from several realizations for the editors in there had been no comprehensive assessments 1997. The first was an awareness that a large of whether these designated areas had been ef- body of biological research from major regions fective in protecting biodiversity. of Costa Rica was available in the literature. In We decided to focus on the seasonal dry for- particular, extensive research had been carried est in the northwestern region of the country out on the lowland Atlantic wet forest, the mid- because it lacked a review comparable to that of dle elevation cloud forest (1,400–1,800 m), and the lowland wet and cloud forests of Costa Rica. the lowland Pacific dry forest over a period of Furthermore, this type of forest was rapidly dis- about 30 years. Further, the wet forest had just appearing, in large part because it was so easily received a comprehensive biological review by converted to agriculture. Daniel Janzen estimated McDade et al. (1994), and a biological review of that only 2 percent of the original Middle Amer- the cloud forest was in progress (see later in this ican tropical dry forest remained. preface). Second, from the 1980s onward, many Building on extensive biological knowledge biologists and nonbiologists made enormous and modern trends for conserving biodiversity, human and financial investments to protect bio- we determined that the book should address diversity in all parts of the country, using mod- three main questions: What do we know about ern conservation approaches and methods. De- the biodiversity and status of the most promi- spite these efforts, little attention had been paid nent groups of plants and animals in the dry to assessing effectiveness of their work. Finally, forest? What have we learned biologically, so- in the more than 30 years that had passed since cioeconomically, and politically about conserv- the first national parks and reserves were estab- ing these specific groups? What do we need to vii frankie front 8/20/03 7:44 PM Page viii consider and do in the future to ensure im- In the current volume we chose to focus on proved conservation and protection of all bio- the lowland seasonal dry-forest region of Costa diversity in the dry forest, as well as other major Rica, which includes several related life zones, regions of the country? according to Holdridge et al. (1971). The book Several major publications have influenced represents the first effort to treat comprehen- our thinking in developing the conceptual goals sively the findings from a wide variety of plant for the book. One of the first workers to investi- and animal biologists investigating a highly gate Costa Rican life zones was Leslie R. Hold- seasonal tropical environment. We were also in- ridge of the Tropical Science Center in Costa terested in whether modern principles of con- Rica. Over a period of several years, he and his servation biology had been put into practice to colleagues intensively studied many plant for- study and conserve dry-forest biodiversity. Thus, mations and their environmental determinants. we asked the biological contributors to assess This work established an ecological foundation the status of the particular taxonomic groups that eventually led to a lengthy treatise on the life they studied and, where applicable, to generalize zone system (Holdridge et al. 1971). Daniel H. for the entire country. In several cases, this re- Janzen (1983) and numerous invited colleagues quest took some authors into other Costa Rican prepared an overview of the natural history of life zones and beyond into adjacent countries the country. His classic 1983 volume and its later of Middle America. Further, because biodiversity Spanish translation (1991) also provided exten- conservation by definition encompasses more sive lists of the plant and animal diversity found than just biology, we also invited several con- in several selected life zones countrywide. tributors to present socioeconomic, policy, legal, More recently, in the 1990s, large scientific and political perspectives on biodiversity conser- publications focused on compiling ecological vation to provide the important social contexts. papers from major life zones that have received Finally, all authors were asked to offer their per- extensive study. For example, McDade et al. sonal recommendations on future directions, (1994) presented a large series of edited papers policies, and actions to better conserve and pro- on the ecology and natural history of a lowland tect biodiversity. These recommendations form Atlantic wet forest, La Selva. This is the site the final synthesis chapter. where researchers associated with the Organi- The book begins with an introductory chap- zation for Tropical Studies first began to study ter, and the chapters that follow are divided into the biota intensively, in 1968; research at La Selva two parts. The final chapter presents conclu- has continued since that time. Nadkarni and sions and recommendations. Chapter 1 (“In- Wheelwright (2000) published a dozen edited troduction”) is concerned with the physical, bio- papers on the natural history and ecology of the logical, human, and conservation environment Monteverde Cloud Forest and its several life of the dry forest.