Alaska’s Changing Boreal Forest F. Stuart Chapin III, et al., Editors OXFORD UNIVERSITY PRESS ALASKA’S CHANGING BOREAL FOREST LONG-TERM ECOLOGICAL RESEARCH NETWORK SERIES LTER Publications Committee Grassland Dynamics: Long-Term Ecological Research in Tallgrass Prairie Edited by Alan K. Knapp, John M. Briggs, David C. Hartnett, and Scott L. Collins Standard Soil Methods for Long-Term Ecological Research Edited by G. Philip Robertson, David C. Coleman, Caroline S. Bledsoe, and Phillip Sollins Structure and Function of an Alpine Ecosystem: Niwot Ridge, Colorado Edited by William D. Bowman and Timothy R. Seastedt Climate Variability and Ecosystem Response in Long-Term Ecological Research Sites Edited by David Greenland, Douglas G. Goodin, and Raymond C. Smith Biodiversity in Drylands: Toward a Unified Framework Edited by 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 Edited by John J. Magnuson, Timothy K. Kratz, and Barbara J. Benson Alaska’s Changing Boreal Forest Edited by 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 Edited by Kris M. Havstad, Laura F. Huenneke, and William H. Schlesinger ALASKA’S CHANGING BOREAL FOREST Edited by F. Stuart Chapin III Mark W. Oswood Keith Van Cleve Leslie A. Viereck David L. Verbyla Illustration Editor Melissa C. Chapin 1 2006 3 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 © 2006 by Oxford University Press, Inc. 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 Alaska’s changing boreal forest / edited by F. Stuart Chapin III . . . [et al.] ; illustration editor, Melissa C. Chapin. p. cm.—(Long-Term Ecological Research Network series) ISBN-10: 0-19-515431-2 ISBN-13: 978-0-19-515431-3 1. Taiga ecology—Alaska. I. Chapin, F. Stuart, III. II. Series. QH105.A4A743 2005 577.3'7'09798—dc22 2004030988 9 8 7 6 5 4 3 2 1 Printed in the United States of America on acid-free paper Preface The boreal forest is the northern-most forested biome, whose organisms and dy- namics have been shaped by low temperature and other features of a high-latitude environment. The Alaskan boreal forest is now warming as rapidly as any place on Earth, providing an unprecedented opportunity to examine a biome as it adjusts to change. This book is written to provide an understanding of the processes that have shaped the development and current dynamics of Alaska’s boreal forest. These processes will constrain its response to future change. Therefore, understanding how Alaskan boreal ecosystems function is essential to guide sound management in a changing climate. We also hope to provide insights for those parts of the world where climate is changing more slowly. The book is intended to provide a synthesis of understanding about Alaska’s boreal forest. It also provides the most complete bibliography currently available for the Alaskan boreal forest and will be a useful guide for those who want to know additional details. It is written in a style that we hope is both scientifically accurate and accessible to general readers who have a curiosity about the North. The first part of the book provides a geographic and historical context for un- derstanding the boreal forest. In the second part, we describe the dynamics of plant and animal communities that inhabit this forest. The third part of the book delves into the biogeochemical processes that link these organisms. The fourth part of the book explores landscape phenomena that operate at larger temporal and spatial scales and integrates the processes described in earlier sections. We are grateful to our co-investigators for the team effort that made this synthe- sis possible. We also thank the many graduate and undergraduate students, post- doctoral fellows, technicians, and colleagues who participated in the research that is synthesized in this book. We particularly thank the site managers (Phyllis Adams, vi Preface Wendy Davis, and Jamie Hollingsworth) and data managers (Darrell Blodgett, Phyllis Adams, Scott Miller, and Brian Riordan), whose dedication and hard work have kept the Bonanza Creek LTER site functioning over the long term. We also recognize the inspiration provided by Keith Van Cleve, Les Viereck, and Ted Dyrness, who initiated the Bonanza Creek LTER program and provided much of its current intellectual focus. We thank the National Science Foundation Long-Term Ecological Research program and the USDA Forest Service Pacific Northwest Experiment Station, which provided the major funding for this research. Finally, we thank the Alaskan boreal forest itself for providing such a magnificent place to work. Contents Contributors xi Part I. Alaska’s Past and Present Environment 1. The Conceptual Basis of LTER Studies in the Alaskan Boreal Forest 3 F. Stuart Chapin III, John Yarie, Keith Van Cleve, and Leslie A. Viereck 2. Regional Overview of Interior Alaska 12 James E. Begét, David Stone, and David L. Verbyla 3. State Factor Control of Soil Formation in Interior Alaska 21 Chien-Lu Ping, Richard D. Boone, Marcus H. Clark, Edmond C. Packee, and David K. Swanson 4. Climate and Permafrost Dynamics of the Alaskan Boreal Forest 39 Larry D. Hinzman, Leslie A. Viereck, Phyllis C. Adams, Vladimir E. Romanovsky, and Kenji Yoshikawa 5. Holocene Development of the Alaskan Boreal Forest 62 Andrea H. Lloyd, Mary E. Edwards, Bruce P. Finney, Jason A. Lynch, Valerie Barber, and Nancy H. Bigelow viii Contents Part II. Forest Dynamics 6. Floristic Diversity and Vegetation Distribution in the Alaskan Boreal Forest 81 F. Stuart Chapin III, Teresa Hollingsworth, David F. Murray, Leslie A. Viereck, and Marilyn D. Walker 7. Successional Processes in the Alaskan Boreal Forest 100 F. Stuart Chapin III, Leslie A. Viereck, Phyllis C. Adams, Keith Van Cleve, Christopher L. Fastie, Robert A. Ott, Daniel Mann, and Jill F. Johnstone 8. Mammalian Herbivore Population Dynamics in the Alaskan Boreal Forest 121 Eric Rexstad and Knut Kielland 9. Dynamics of Phytophagous Insects and Their Pathogens in Alaskan Boreal Forests 133 Richard A. Werner, Kenneth F. Raffa, and Barbara L. Illman 10. Running Waters of the Alaskan Boreal Forest 147 Mark W. Oswood, Nicholas F. Hughes, and Alexander M. Milner Part III. Ecosystem Dynamics 11. Controls over Forest Production in Interior Alaska 171 John Yarie and Keith Van Cleve 12. The Role of Fine Roots in the Functioning of Alaskan Boreal Forests 189 Roger W. Ruess, Ronald L. Hendrick, Jason G. Vogel, and Bjartmar Sveinbjörnsson 13. Mammalian Herbivory, Ecosystem Engineering, and Ecological Cascades in Alaskan Boreal Forests 211 Knut Kielland, John P. Bryant, and Roger W. Ruess 14. Microbial Processes in the Alaskan Boreal Forest 227 Joshua P. Schimel and F. Stuart Chapin III 15. Patterns of Biogeochemistry in Alaskan Boreal Forests 241 David W. Valentine, Knut Kielland, F. Stuart Chapin III, A. David McGuire, and Keith Van Cleve Contents ix Part IV. Changing Regional Processes 16. Watershed Hydrology and Chemistry in the Alaskan Boreal Forest: The Central Role of Permafrost 269 Larry D. Hinzman, W. Robert Bolton, Kevin C. Petrone, Jeremy B. Jones, and Phyllis C. Adams 17. Fire Trends in the Alaskan Boreal Forest 285 Eric S. Kasischke, T. Scott Rupp, and David L. Verbyla 18. Timber Harvest in Interior Alaska 302 Tricia L. Wurtz, Robert A. Ott, and John C. Maisch 19. Climate Feedbacks in the Alaskan Boreal Forest 309 A. David McGuire and F. Stuart Chapin III 20. Communication of Alaskan Boreal Science with Broader Communities 323 Elena B. Sparrow, Janice C. Dawe, and F. Stuart Chapin III 21. Summary and Synthesis: Past and Future Changes in the Alaskan Boreal Forest 332 F. Stuart Chapin III, A. David McGuire, Roger W. Ruess, Marilyn W. Walker, Richard D. Boone, Mary E. Edwards, Bruce P. Finney, Larry D. Hinzman, Jeremy B. Jones, Glenn P. Juday, Eric S. Kasischke, Knut Kielland, Andrea H. Lloyd, Mark W. Oswood, Chien-Lu Ping, Eric Rexstad, Vladimir E. Romanovsky, Joshua P. Schimel, Elena B. Sparrow, Bjartmar Sveinbjörnsson, David W. Valentine, Keith Van Cleve, David L. Verbyla, Leslie A. Viereck, Richard A. Werner, Tricia L. Wurtz, and John Yarie Index 339
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