Tropical Forest Ecology: A View from Barro Colorado Island

262 Pages·1999·28.77 MB·English

by  Egbert Giles Leigh Jr.

TROPICAL FOREST ECOLOGY
This page intentionally left blank
TROPICAL FOREST ECOLOGY
A VIEW FROM BARRO COLORADO ISLAND
EGBERT GILES LEIGH, JR.
New York Oxford
Oxford University Press
1999
Oxford University Press
Oxford New York
Athens Auckland Bangkok Bogota Buenos Aires Calcutta
Cape Town Chennai Dar os Salaam Delhi Florence Hong Kong Istanbul
Karachi Kuala Lumpur Madrid Melbourne Mexico City Mmnbai
Nairobi Paris Sao Paulo Singapore Taipei Tokyo Toronto Warsaw
and associated companies in
Berlin Ibadan
Copyright © 1999 by Oxford University Press, Inc.
Published by Oxford University Press, Inc.
198 Madison Avenue, New York, New York 1001B
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-Publicalion Data
Leigh, Egbert Giles.
Tropical forest ecology : a view from
Barro Colorado Island / by
Egbert Giles Leigh, Jr.
p. cm.
Includes bibliographical references and index.
ISBN 0-19-509602-9; 0-19-509603-7 (pbk.)
1. Forest ecology—Panama—Barro Colorado Island. 2. Barro
Colorado Island (Panama) I. Title.
QH108.P3L45 1999
577.34'097287'5—dc21 97-13770
1 3 5 7 9 8 6 42
Printed in the United States of America
on acid-free paper
This book is dedicated to the memory of Martin
Moynihan (1928-1096), the founding director of
the Smithsonian Tropical Research Institute, in
thanks For giving me the opportunity to become
a tropical biologist, arranging my acquaintance
with a splendid selection of rainforests, and pro-
viding me with such a superb setting in which
to practice my trade,
This page intentionally left blank
Preface
Tropical forest has provided me with a large share of that Polanyi (1958) has shown how dangerous it is to try to cut
experience of beauty without which any life would be sadly the person out of the scientist. Instead, I wish to interest
incomplete. Indeed, I confess to being a creature of my readers of goodwill in tropical forests and what can be
memories of these experiences—the late afternoon sun learned from them. I find this task rather urgent, as I am
lending color to the tree trunks of Barro Colorado during convinced that the road to a proper understanding of ecol-
walks early in the dry season; sunrise over the misty for- ogy and evolution begins and ends in the tropics.
est on that island after a night of rain, •waiting for the howler In 1967, the founding director of the Smithsonian Tropi-
monkeys to howl and be counted; the eerie, rather musical cal Research Institute (STRI) in Panama, Martin Moynihan,
whoops of babakotos, Madagascar's largest living lemurs, was encouraging studies to learn how representative the
as they jumped gracefully from tree to tree in the rain- ecology of Barro Colorado's forest and the behavior of its
forest near Perinet; "conversations" with lemurs near a animals were of tropical forests in general. As a young theo-
ridgetop camp overlooking what was then unbroken rain- retical biologist, I felt that comparing ecological commu-
forest sloping to the sea in northeastern Madagascar; the nities in different biogeographic realms, with different
pastel clouds shifting about over the Bay of Antongil at evolutionary histories, would be a good way to acquaint
sunset as I was watching from a cloud forest mountaintop myself with the level of detail theoretical prediction ought
on this same Malagasy peninsula; the redwood-like gran- to strive for. Moynihan engaged me to compare the struc-
deur of the enormous dipterocarps, Shorea curtisii, with ture and physiognomy of tropical forests around the world
elegant crowns full of small, silvery leaves, crowning the and he sent me and my wife, the former Elizabeth Hodgson,
rainforest on the flank of a ridge at Ulu Gombak, northeast on two trips around the world, in 1968 and 1970, to do this.
of Kuala Lumpur; eating my first durian in the heath for- He also appointed me to the STRI staff in 1969, thus shelter-
est atop Gunong Ulu Kali, less than an hour's drive from ing me from the transformations of American universities,
Kuala Lumpur; and many others. formerly citadels of thought, into organizations •where,
This book is not a record of my memories—after all, I nowadays, no one has a moment in which to stop and
have not Proust's skill. Rather, it is an attempt to make think. The problem of the similarities and differences
sense of them. It is an account of my understanding of tropi- among tropical forests of different regions continues to
cal forest, as it has developed through my own experience, preoccupy me, This book is a belated attempt to answer
studies of many of my friends on Barro Colorado and else- Moynihan's question. As I owe my career, my involvement
where, and my reading. Thus this book is inevitably a very in tropical biology, the intellectual atmosphere which pre-
personal one. I will not try to review the literature about vails on Barro Colorado and elsewhere at STRI, and per-
tropical forest. Still less do I claim to present the current haps even my marriage, as well as this book's central ques-
consensus among tropical biologists about the organizing tion, to Martin Moynihan, I dedicate this book to him.
principles of tropical forest—I don't think there is any such This book owes much to many others. A. G. Fischer
consensus, and if there were, my experience of scientific introduced me to the fossil record and to ecology as a tool
fads and bandwagons would lead me to avoid it like the for making sense of that record. Robert Stallard introduced
plague. I have no desire to lull my readers to sleep with me to the processes of weathering and soil formation. Rob-
the deadly ether of soi-disant impersonal objectivity: ert MacArthur and James Crow introduced me to Ronald
VIII PREFACE
Fisher's (1930) Genetical Theory of Natural Selection. these trips also allowed me to see several of the many dif-
MacArthur and Crow helped show me the relevance of the ferent types of forest in south India.
common interest among a genome's genes in distinguish- Many people have supplied me with reprints and un-
ing between the selfish advantage of individual alleles and published data. I must single out Robin Foster and Rich-
the fitness of genotypes. MacArthur also taught me how ard Condit, and their data analysts, Una Smith and Suzanne
to think about biotic diversity. This book also owes an Loo de Lao, for the abundant data they have provided me
enormous debt to E. J. H. Corner. I never met him, but his on the 50-ha Forest Dynamics Plot on Barro Colorado and
papers, particularly his essay (Corner 1954) on the evo- similar plots maintained by the Center for Tropical Forest
lution of tropical forest, his book (Corner 1964), The Life Science elsewhere in the world. Similarly, I am grateful to
of Plants, and his monograph (Corner 1967) on the figs of Donald Windsor, S. Joseph Wright, and Steven Paton for
the Solomon Islands, have given me abundant food for supplying data and analyses from monitoring records of
thought. Corner brings out, as no one else could, the idea the Smithsonian's Environmental Sciences Program.
of tropical forest as a mutualism; moreover, no one else S. J. Wright and G. J. Vermeij read the whole manuscript
can communicate so effectively the beauty and mystery with care and called my attention to errors, omissions, and
of tropical forest. Finally, Russell Lande, Montgomery obscurely written passages. David King and Hubert Herz
Slatkin, and David Sloan Wilson, among others, showed read chapters 5 and 6, and Richard Condit read chapter 8;
me how to reconcile Fisher's approach with Corner's. they all warned me of other ditches I was falling into. The
My wife, Elizabeth, helped me with most of the field- errors that remain, of course, are my own, but, without their
work reported in this book. Robin Foster introduced me to help, there would have been many more.
the plants of Barro Colorado and showed me how popula- I must also thank a splendidly various company of art-
tions of vertebrate herbivores were limited by seasonal ists for their efforts to capture one or another aspect of tropi-
shortage of fruit and new leaves. Marcel and Annette cal nature, among them, Alex Murawski, Daniel Glanz,
Hladik, who were predoctoral fellows on Barro Colorado George Angehr, Judith Gradwohl, Arlee Montalvo, Lynn
when I first went there, gave me a 12-year subscription to Siri Kimsey, Scott Gross, Gerardo Ravassa, Karen Kraeger,
La Terre et la Vie, and otherwise kept me in touch with Roxanne Trapp, Marshall Hasbrouck, Hayro Cunampio,
French work in Africa and South America, a very precious Francesco Gattesco, Donna Conlon, Dorsett W. Trapnell,
gift. In 1991, they involved me in a UNESCO conference and Deborah Miriam Kaspari. Steven Paton, Robert Stal-
on rainforest peoples, food, and nutrition (Hladik et al. lard, and David Kinner supplied the maps of Barro Colo-
1993), which showed me the many lessons to be learned rado Island and the Barro Colorado Nature Monument.
from these people, not least the elementary lesson, forgot- I also thank a host of librarians, especially Carol Jopling,
ten by the intellectual elites of the west, that we do not live Sylvia Churgin, Tina Lesnick, Vielka Chang-yau, and
by bread alone. The Hladiks also provided me the oppor- Angel Aguirre, for procuring copies of all sorts of obscure
tunity to assemble, summarize, and publish my thinking papers. I thank the American Philosophical Society and
on the role of mutualism in evolution: this paper (Leigh Princeton University for help toward my first trip around
and Rowell 1995) was in many ways a trial run for chapter the world, and the Smithsonian Research Opportunities
9. Allen Herre (1996) has shown how a single mutualism, Fund, its predecessor, and the Smithsonian Tropical Re-
that between fig trees and their pollinating wasps, can search Institute for other financial support.
shape not only the biology of fig trees, but the ecological I also wish to express my gratitude to the Panamanian
characteristics of a tropical forest. government for its hospitality to the Smithsonian Tropi-
Indeed, this story spreads far beyond Barro Colorado. I cal Research Institute, and therefore, to me. In an age where
am indebted to Benjamin Stone, formerly of the Univer- many countries view scientific research as another form
sity of Malaya, for introducing me to durians and diptero- of exploitation, to be strictly regulated and controlled, it
carps, cloud forests and Leptospermum heaths, in the is a pleasure to acknowledge that in Panama the honor of
Malay Peninsula. His recent death was almost as severe a knowledge for the sake of knowing, so characteristic of the
blow to tropical botany as the death of Al Gentry. Alison Enlightenment, is still very much alive.
Jolly, Vololoniaina Jeannoda, Rakotonirina Bonja, and Jean Finally, I thank the current director of the Smithsonian
Prosper Abraham have all helped acquaint me with the Tropical Research Institute, Ira Rubinoff, for his continued
marvelous diversity of plants and habitats in Madagascar. support and encouragement. My style of work is out of fash-
The Smithsonian Foreign Currency Program, which fi- ion, and I bring no money to the institute, yet he has left me
nanced our initial trips around the world, made possible the freedom to do my work in peace, with ample encour-
subsequent visits to India. Twice, in 1985 and 1995, Madhav agement and the necessary financial support. Meanwhile, he
Gadgil, Raghavendra Gadagkar, and R. Sukumar invited me has managed to guide this institute through an endless se-
to the Centre for Ecological Science at the Indian Institute ries of political minefields without endangering its commit-
of Science in Bangalore to lecture, first on why there are so ment to basic research. He has diversified its activities; and
many kinds of tropical trees, and later on the ecology of Barro thanks to him, I have the opportunity to hobnob with and
Colorado Island. These lectures were marvelous opportu- learn from plant physiologists, anthropologists, archaeolo-
nities to collect and organize my thoughts for this book; gists, palynologists, and even, horror of horrors, the occa-
PREFACE IX
sional molecular biologist, as well as the ethologists, ecolo- Corner, )i. J. H. 1967. Ficus in the Solomon Islands and its bear-
ing on the post-Jurassic history of Melanesia. Philosophical
gists, and naturalists who first established the patterns of basic
Transactions of the Royal Society of London, B 253: 23-159.
research at STRI. This book has benefitted from all of them.
Fisher, K. A. 1930. The Genetical Theory of Natural Selection.
Clarendon Press, Oxford.
Barro Colorado Island E. G. L., Jr.
Horn-!, E. A. 1996. An overview of studies on a community of
Feast of the Conversion of St. Paul Panamanian figs. Journal of Biogeography 23: 593-607.
1997 Hladik, C. M., A. Hladik, O. F. Linares, H. Pagezy, A. Semple,
and M. Hadley (eds). 1993. Tropical Forests, People and
Food. UNESCO, Paris, and Parthenon Publishing, Park Ridge,
References
NJ.
Corner, E. J. H. 1954. The evolution of tropical forest, pp. 34—46. Leigh, E. G. Jr., and T. E. Rowell. 1995. The evolution of mutual-
In J. Huxley, A. C. Hardy, and E. B. Ford, eds., Evolution as ism and other forms of harmony at various levels of biologi-
a Process. George Allen and Unwin, London. cal organization. Ecologie 26: 131-158.
Comer, E. J. H. 1964. The Life of Plants. WorJd Publishing, Cleve- Polauyi, M. 1958. Personal Knowledge. University of Chicago
land, OH. Press, Chicago.