Principios De Ecología

F. Stuart Chapin III Pamela A. Matson Harold A. Mooney

Principles of Terrestrial Ecosystem Ecology
Illustrated by Melissa C. Chapin

With 199 Illustrations

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F. Stuart Chapin III Institute of Arctic Biology University of Alaska Fairbanks, AK 99775 USA terry.chapin@uaf.edu Harold A. Mooney Department of Biological Sciences Herrin Hall, MC 5020 Stanford University Stanford, CA94305-5020 USA hmooney@jasper.stanford.edu

Pamela A. Matson Department of Geological and Environmental Science School of Earth Sciences Green 355 Stanford University Stanford, CA 94305-2115 USA matson@pangea.stanford.edu

Cover illustration: Waterfall and forests on Valean Poas in Costa Rica. Photograph by Peter Vitousek.

Library of Congress Cataloging-in-Publication Data Chapin, F. Stuart(Francis Stuart), III. Principles of terrestrial ecosystem ecology / F. Stuart Chapin III, Pamela A. Matson, Harold A. Mooney. p. cm. Includes bibliographical references (p. ) ISBN 0-387-95439-2 (hc :alk. paper)—ISBN 0-387-95443-0 (sc :alk. paper) 1. Ecology. 2. Biogeochemical cycles. 3. Biological systems. I. Matson, P. A. (Pamela A.) II. Mooney, Harold A. III. Title. QH541 .C3595 2002 577¢.14—dc212002017654 ISBN 0-387-95439-2 ISBN 0-387-95443-0 (hardcover) (softcover) Printed on acid-free paper.

© 2002 Springer-Verlag New York, Inc. All rights reserved. This work may not be translated or copied in whole or in part without the written permission of the publisher (Springer-Verlag New York, Inc., 175 Fifth Avenue, New York, NY 10010, USA), except for brief excerpts in connection with reviewsor scholarly analysis. Use in connection with any form of information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed is forbidden. The use in this publication of trade names, trademarks, service marks, and similar terms, even if they are not identified as such, is not to be taken as an expression of opinionas to whether or not they are subject to proprietary rights. Printed in the United States of America. 9 8 7 6 5 4 3 2 1 www.springer-ny.com Springer-Verlag New York Berlin Heidelberg A member of BertelsmannSpringer Science+Business Media GmbH SPIN 10866301 (hardcover) SPIN 10866759 (softcover)

Preface

Human activities are affecting the global environment in myriad ways, with numerous directand indirect effects on ecosystems. The climate and atmospheric composition of Earth are changing rapidly. Humans have directly modified half of the ice-free terrestrial surface and use 40% of terrestrial production. Our actions are causing the sixth major extinction event in the history of life on Earth and are radically modifying the interactions among forests, fields, streams, and oceans. Thisbook was written to provide a conceptual basis for understanding terrestrial ecosystem processes and their sensitivity to environmental and biotic changes. We believe that an understanding of how ecosystems operate and change must underlie our analysis of both the consequences and the mitigation of human-caused changes. This book is intended to introduce the science of ecosystem ecology to advancedundergraduate students, beginning graduate students, and practicing scientists from a wide array of disciplines. We also provide access to some of the rapidly expanding literature in the many disciplines that contribute to ecosystem understanding. The first part of the book provides the context for understanding ecosystem ecology. We introduce the science of ecosystem ecology and place it in thecontext of other components of the Earth System—the atmosphere, ocean, climate and geological systems. We show how these components affect ecosystem processes and contribute to the global variation in terrestrial ecosystem structure and processes. In the second part of the book, we consider the mechanisms by which terrestrial ecosystems function and focus on the flow of water and energy and the...