OUP UNCORRECTED PROOF A CAribbeAn Forest Tapestry BROKAW-halftitle1-PageProof i January 12, 2012 8:24 PM OUP UNCORRECTED PROOF LONG-TERM ECOLOGICAL RESEARCH NETWORK SERIES LTER Publications Committee Grassland Dynamics: Long-Term Ecological Research in Tallgrass Prairie Editors: Alan K. Knapp, John M. Briggs, David C. Hartnett, and Scott L. Collins Standard Soil Methods for Long-Term Ecological Research Editors: G. Philip Robertson, David C. Coleman, Caroline S. Bledsoe, and Phillip Sollins Structure and Function of an Alpine Ecosystem: Niwot Ridge, Colorado Editors: William D. Bowman and Timothy R. Seastedt Climate Variability and Ecosystem Response at Long-Term Ecological Sites Editors: David Greenland, Douglas G. Goodin, and Raymond C. Smith Biodiversity in Drylands: Toward a Unified Framework Editors: Moshe Shachak, James R. Gosz, Steward T.A. Pickett, and Avi Perevolotsky Long-Term Dynamics of Lakes in the Landscape: Long-Term Ecological Research on North Temperate Lakes Editors: John J. Magnuson, Timothy K. Kratz, and Barbara J. Benson Alaska’s Changing Boreal Forest Editors: F. Stuart Chapin III, Mark W. Oswood, Keith Van Cleve, Leslie A. Viereck, and David L. Verbyla Structure and Function of a Chihuahuan Desert Ecosystem: The Jornada Basin Long-Term Ecological Research Site Editors: Kris M. Havstad, Laura F. Huenneke, and William H. Schlesinger Principles and Standards for Measuring Net Primary Production in Long-Term Ecological Studies Editors: Timothy J. Fahey and Alan K. Knapp Agrarian Landscapes in Transition: Comparisons of Long-Term Ecological and Cultural Change Editors: Charles L. Redman and David R. Foster Ecology of the Shortgrass Steppe: A Long-Term Perspective Editors: William K. Lauenroth and Ingrid C. Burke BROKAW-seriespage-PageProof ii January 12, 2012 6:33 PM OUP UNCORRECTED PROOF A Caribbean Forest Tapestry The Multidimensional Nature of Disturbance and Response Edited by nicholas brokaw, todd a. crowl, ariel e. lugo, william h. mcdowell, frederick n. scatena, robert b. waide, and michael r. willig BROKAW-titlepage-PageProof iii January 12, 2012 6:46 PM OUP UNCORRECTED PROOF Oxford University Press, Inc., publishes works that further Oxford University’s objective of excellence in research, scholarship, and education. Oxford New York Auckland Cape Town Dar es Salaam Hong Kong Karachi Kuala Lumpur Madrid Melbourne Mexico City Nairobi New Delhi Shanghai Taipei Toronto With offices in Argentina Austria Brazil Chile Czech Republic France Greece Guatemala Hungary Italy Japan Poland Portugal Singapore South Korea Switzerland Thailand Turkey Ukraine Vietnam Copyright © 2012 by Oxford University Press Published by Oxford University Press, Inc. 198 Madison Avenue, New York, New York 10016 www.oup.com Oxford is a registered trademark of Oxford University Press All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise, without the prior permission of Oxford University Press. Library of Congress Cataloging-in-Publication Data A Caribbean forest tapestry : the multidimensional nature of disturbance and response / edited by Nicholas Brokaw . [et al.]. p. cm. Includes bibliographical references and index. ISBN 978-0-19-533469-2 (hardcover : acid-free paper) 1. Forest ecology—Puerto Rico—Luquillo Mountains. 2. Forest ecology—Caribbean Area. 3. Forest ecology—Tropics. 4. Ecological disturbances— Puerto Rico—Luquillo Mountains. 5. Adaptation (Biology)—Puerto Rico—Luquillo Mountains. 6. Biotic communities—Puerto Rico—Luquillo Mountains. 7. Forest management—Puerto Rico—Luquillo Mountains. 8. Forest conservation—Puerto Rico—Luquillo Mountains. 9. Luquillo Mountains (P.R.)— Environmental conditions. I. Brokaw, Nicholas V. L. QH109.P6C37 2012 577.3097295—dc23 2011035972 1 3 5 7 9 8 6 4 2 Printed in the United States of America on acid-free paper BROKAW-copyright-PageProof iv January 13, 2012 3:29 PM OUP UNCORRECTED PROOF Dedication This book is dedicated to two influential individuals who pioneered the fields of tropical forestry and tropical ecosys- tem ecology. Frank H. Wadsworth, a forester, helped to institutionalize tropical forestry in the Neotropics by orga- nizing forest services and training foresters in several coun- tries, establishing the regional research journal Caribbean Forester, and helping in the development of the Latin American Forestry Commission of the Food and Agricul- ture Organization some 50 years ago. Howard T. Odum, an ecologist, revolutionized the study and interpretation of tropical forests with the application of thermodynamics to ecosystem analysis and the use of large-scale studies such as the giant cylinder to study the metabolism of tropical forests. The approach and focus of this book rests on the shoulders of these two exceptional scientists who dedi- cated a considerable portion of their careers to under- standing the functioning of the ecosystems of the Luquillo Mountains (LM). Wadsworth was transferred to Puerto Rico by the U.S. Forest Service in 1942, and Odum first visited the LM in 1944 to learn tropical meteorology as a 2nd Lieutenant of the U.S. Army. Among the many tropical forestry is- sues that Wadsworth addressed throughout his career in Puerto Rico, three are immediately relevant to this book. BROKAW-dedication-PageProof v January 13, 2012 3:22 PM OUP UNCORRECTED PROOF He collaborated with Elbert L. Little, Jr., and Roy O. Woodbury in the description of all tree species of Puerto Rico; he established what is now the oldest network of tree-growth plots in the Neotropics; and he manipulated forest basal areas in order to test ideas about tropical forest management. Wadsworth was interested in under- standing tree growth in the tropics and in developing prescriptions for their management. Odum returned to the LM in 1957 on a Rockefeller grant, and with colleagues such as Frank B. Golley, he began studying the metabolism of mangroves and wet tropical forests, including their biomass and carbon se- questration. By 1962, Odum had developed whole ecosys- tem models with data collected in the LM and an energy language that changed the way ecologists analyzed trop- ical forests. During these visits, he and Wadsworth dis- cussed fundamental issues of tropical forest management, as is evident from the following quotation from Odum (1962:66): The preliminary calculations [of the organic matter flux] pro- vide a possible solution to one question troubling Dr. Frank Wadsworth and associates, tropical foresters managing this forest. The growth rate of trees measured over 20 years has been small, 0.05 to 0.12 inches per year. The dominant trees are several hundred years old. Is this slow growth due to lack of light, lack of nutrients, or inadequate photosynthesis for other reasons? The calculation of respiration as 9 gm2 day−1 due to leaves and 5.8 due to the soil, root, and litter indi- cates very little production is left for any net growth with most of it being used to sustain leaf and soil activity. The apparent reason for slow growth is thus not any inhibition of gross photosynthesis, but the full development of the eco- system structure requiring most of the production for respi- ratory maintenance. Odum’s work in Sabana evolved into the Rain Forest Ra- diation Study, funded by the U.S. Atomic Energy Commis- sion and hosted by the University of Puerto Rico (UPR). This study is recognized as the first example of a “big science” ecosystem project in the tropics, a harbinger of the Long Term Ecological Research (LTER) program. The outcomes of the collaboration between Wadsworth and Odum were many but are highlighted by the allocation of 180 acres of National Forest lands for exclusive research use by the UPR; the use of Forest Service facilities in support of UPR research activities, including the library and headquarters of what BROKAW-dedication-PageProof vi January 13, 2012 3:22 PM OUP UNCORRECTED PROOF was then Institute of Tropical Forestry; and the sharing of information and ideas about tropical forests. These out- comes and the research infrastructure created by both Wadsworth and Odum became the backbone of the Luquillo LTER program, the results of which are summa- rized in this book. However, as important as the research infrastructure has been to the LTER program, it was the ideas that emerged from the friendship between Wadsworth and Odum that constitutes their greatest legacy, a legacy that is so evident in this book. This intellectual legacy was cemented in their independent minds, their focus on experimentation at the ecosystem level, and their openness to innovation. They both understood the tropical forest as a system without ignoring the importance of its parts. It is not an accident that the Luquillo LTER has been successful in the integra- tion of population and ecosystem ecology. Both Wadsworth and Odum successfully supported population ecology research while also maintaining a whole-system perspec- tive and fomenting whole-system research. They both had a worldview and understood that science is a vehicle for helping resolve conservation issues and for addressing human needs. Such ideas are evident in the books in which they independently culminated their career experiences in Puerto Rico (Odum 1971; Wadsworth 1997). This book extends their points of view and celebrates the intellectual synergy that they displayed between 1963 and 1989. Literature Cited Odum, H. T. 1962. Man and the ecosystem. Pages 57–75 in P. E. Waggoner and J. D. Oving- ton, editors, Proceedings of the Lockwood conference on the suburban forest and ecology. New Haven, CT: Connecticut Agricultural Experiment Station. Odum, H. T. 1971. Environment, power and society. New York: Wiley Interscience. Wadsworth, F. H. 1997. Forest production for tropical America. USDA Forest Service Agri- culture Handbook 710. Washington, DC: USDA Forest Service. BROKAW-dedication-PageProof vii January 13, 2012 3:22 PM OUP UNCORRECTED PROOF Contents Preface xi Acknowledgments xvii Contributor xxi 1 EcologicalParadigmsfortheTropics Old Questions and Continuing Challenges 3 Ariel E. Lugo, Robert B.