Ecological Studies, Vol. 166 Analysis and Synthesis Edited by I.T. Baldwin, Jena, Germany M.M. Caldwell, Logan, USA G. Heldmaier, Marburg, Germany O.L. Lange, Wiirzburg, Germany H.A. Mooney, Stanford, USA E.-D. Schulze, Jena, Germany U. Sommer, Kie1, Germany Ecological Studies Volumes published since 1996 are listed at the end of this book. Springer Science+Business Media, LLC Paul J. Hanson Stan D. Wullschleger Editors North American Temperate Deciduous Forest Responses to Changing Precipitation Regimes Foreword by Jerry W. Elwood With 129 Figures, 4 in Full Color , Springer Paul J. Hanson Stan D. Wullschleger Environmental Sciences Division Environmental Sciences Division Oak Ridge National Laboratory Oak Ridge National Laboratory Bethel Valley Road, Building 1059 Bethel Valley Road, Building 1059 P.O. Box 2008 P.O. Box 2008 Oak Ridge, TN 37831-6422 Oak Ridge, TN 37831-6422 USA USA [email protected] [email protected] Cover illustration: Designed by LeJean Hardin of the Oak Ridge National Laboratory's Creative Media Services Division, Oak Ridge, Tennessee. Library of Congress Cataloging-in-Publication Data North America temperate deciduous forest responses to changing precipitation regimes / editors, Paul 1. Hanson, Stan D. Wullschleger. p. cm. - (Ecological studies ; 166) Includes bibliographical references (p. ). ISBN 978-1-4612-6506-1 ISBN 978-1-4613-0021-2 (eBook) DOI 10.1007/978-1-4613-0021-2 1. Rain and rainfall-North America. 2. Forest ecology-North America. 3. Precipitation variability-North America. 4. Climatic changes-North America. I. Hanson, Paul J. (Paul James) 11. Wullschleger, Stan D. III. Ecological studies ; v. 166. SD390.7.R34N672003 577.3'097-dc21 2002044525 ISSN 0070-8356 ISBN 978-1-4612-6506-1 Printed on acid-free paper. © 2003 Springer Science+Business Media New Y ork Originally published by Springer-Verlag New York, Inc. in 2003 Softcover reprint of the hardcover I st edition 2003 All rights reserved. This work may not be translated or copied in whole or in part without the wrilten permission of the publisher (Springer Science+Business Media, LLC), except for brief excerpts in connection with reviews or scholarly analysis. Use in connection with any form of information storage and retrieval, e1ectronic adaptation, computer software, or by similar or dissimilar methodology now known or here after 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 opinion as to whether or not they are subject to proprietary rights. 9 8 7 6 5 4 3 2 1 SPIN 10906231 www.springer-ny.com Foreword Ecologists have long been active in conducting observational studies and exper imental research aimed at better understanding the response of terrestrial ecosys tems to changes in their chemical and physical environment, including changes in climate. Early studies that focused on primary and secondary response of plants to air pollution gave way to studies that dealt with acidic precipitation, that in turn gave way to studies on pressing concerns of global change. Understanding the effects of increases in atmospheric CO concentration, global warming, and 2 regional changes in precipitation offers a unique set of challenges for the scientific community. The interagency United States Global Change Research Program (USGCRP), established in 1989 and authorized by Congress in the Global Change Research Act of 1990, has been an advocate for research in these and other areas. A goal of the USGCRP is to understand and predict the consequences of global environmental change in terrestrial ecosystems. As a participating agency in the USGCRP, the Department of Energy (DOE) shares this goal through our Program for Ecosystem Research (PER) which seeks to improve the scientific basis for predicting and assessing effects of environmental changes on terrestrial ecosystems and their com ponent organisms and processes. Specific goals of PER include understanding (1) how organisms within terrestrial ecosystems respond to environmental change, (2) how these responses are controlled, (3) how the responses and their underlying control mechanisms can be detected and measured, and (4) how responses of or- v vi Foreword ganisms can be integrated across organizational levels from the results of studies of specific research organisms to ecosystems. Given these many objectives, the DOE Office of Biological and Environmental Research in the Office of Science began funding the Throughfall Displacement Experiment (IDE) in 1992. The results were expected to provide an improved mechanistic understanding of how upland mixed deciduous forests respond to altered precipitation amount. The 26 chapters of this book demonstrate that many of our expectations have been fulfilled. They represent an important source of information regarding the resiliency and/or vulnerability of our nation's decidu ous forests to long-term (10 years to date) altered precipitation amount. Results from the TDE are important not only from the perspective of understanding the many mechanisms by which temperate deciduous forests respond to changes in the amount of precipitation, but also provide new information and insights needed for predicting and assessing whole forest responses to possible future changes. Such is the opportunity afforded by long-term studies like the TDE that are con ducted at ecosystem spatial scales. It remains a challenge to incorporate the data and conclusions reported herein into future national assessments of climate change impacts, and make them available for discussion within the policy arena. Comprehensive and long-term data, from the IDE and similar studies, present the scientific communities with a unique opportunity to judge the efficacy of ecosystem models that are ultimately useful for forecasting effects of possible future environmental changes on the structure and functioning of ecosystems. Finally, operation of the IDE since 1992 has required a long-term commitment of resources, and periodic peer-review has been required to support this sustained research. The goal has been to operate such studies, with continuing input and guidance from the scientific community, so that potentially important, but slow to-develop, ecosystem responses have time to express themselves and be detected. As the IDE progresses towards its completion over the next several years, some conclusions will be strengthened but other insights may need to be modified or abandoned in favor of alternative explanations. Such is the scientific process and such is the return that can only come from dedicated investments in long-term studies. Jerry W. Elwood Director, Climate Change Research Division Office of Biological and Environmental Research Office of Science U.S. Department of Energy Preface The current volume is in many respects the next issue in a series of books by Springer-Verlag that summarize fundamental research on functional processes operating in eastern deciduous forested ecosystems. Analysis of Temperate Forest Ecosystems (Reichle 1970) provided a broad description of temperate forest eco systems and their nutrient and hydrologic cycles. That book resulted mainly from research on the Eastern Deciduous Forest Biome as a part of the International Biological Programme. Four books, Biogeochemistry of a Forested Ecosystem (Likens et al. 1977), Pattern and Process in a Forested Ecosystem (Borman and Likens 1979), Forest Hydrology and Ecology at Coweeta (Swank and Crossley 1988), and Analysis of Biogeochemical Cycling Processes in Walker Branch Watershed (Johnson and Van Hook 1989), include long-term studies of watershed scale processes, including forest-management perspectives. Atmospheric Depo sition and Forest Nutrient Cycling (Johnson and Lindberg 1992), motivated by the acid rain issue of the 1980s, includes thorough descriptions of nutrient cycling processes and ozone exposure in a variety of forested ecosystems including south ern and northern hardwood sites. The present book was based on a need to better understand deciduous forested ecosystem responses to potential climatic condi tions of the future. We hope that this work, when combined with its predecessor volumes, will provide important information on mechanisms of eastern deciduous forest plant and ecosystem responses to altered precipitation patterns, and that this will enhance assessments of potential impacts of environmental change on forested ecosystems. vii viii Preface Data from the Walker Branch Throughfall Displacement Experiment (TDE) detailed in this volume has been archived by the Carbon Dioxide Information and Analysis Center (CDIAC) and can be found at the following web address: http:// cdiac.esd.oml.gov/epubs/ndp/ndp078a/ndp078a.html. The archive includes data for site characterization, environment conditions, soil-water content, growth, sea sonal phenology, plant physiology, soil respiration and decomposition, and nu trient cycling necessary for future development and testing of ecosystem models of upland oak deciduous forests. Paul J. Hanson Stan D. Wullschleger References Bonnan FH, Likens GE (1979) Pattern and process in a forested ecosystem. Springer-Verlag, New York. Likens GE, Bonnann FH, Pierce RS, Eaton JS, Johnson NM (1977) Biogeochemistry of a Forested Ecosystem. Springer-Verlag, New York. Johnson DW, Lindberg SE Editors (1992) Atmospheric deposition and forest nutrient cycling. Springer-Verlag, New York. Johnson DW, Van Hook RJ Editors (1989) Analysis of biogeochemical cycling processes in Walker Branch Water shed. Springer-Verlag, New York. Reichle DE Editor (1970) Analysis of temperate forest ecosystems. Springer-Verlag, New York. Swank WT, Crossley DA Jr. Editors(1988) Forest hydrology and ecology at Coweeta. Springer-Verlag, New York. Acknowledgments All research on the Walker Branch Watershed Throughfall Displacement Exper iment (TDE) was sponsored in full or in part by the Office of Science, Biological and Environmental Research (BER) Program, U.S. Department of Energy (DOE) as a part of the Program for Ecosystem Research. Work conducted by Oak Ridge National Laboratory personnel was conducted under contract No. DE-AC05 OOOR22725 with UT-Battelle, LLC. The TDE is on the Oak Ridge National Environmental Research Park. Many people contributed to the completion of this book. Frederick M. O'Hara, Jr., did an excellent job as technical editor of the volume. Not only did he carefully edit each chapter, but in some cases he suggested major revisions of sections. The authors sincerely appreciate his knowledge of science, his humor, and his tenacity in performing the many tasks needed to bring this book to pUblication. We thank the following students, faculty participants, and co-workers for par ticipating in the long-term collection of data used in several chapters of this book: Philip Allen, Steve Armes, James E. Arnett III, Stephanie Bohlman, Bridgette Boudreaux, Mogan Castner, Jason Fults, Kristin Harter, Michael Hayes, David Holt, Joshua Johnson, Sabina Madsen, Howard Marotto, Tracy Misek, Mark Scannell, Kevin Smith, James B. Stringfellow, Todd Tabberer, and Karen Voiles. The authors of Chapter 5 thank Donald E. Todd for the extensive effort in volved in sample collection and Ramie V. Wilkerson for excellent technical support. ix x Acknowledgments Work on Chapter 7 was supported by The University of Tennessee Agricultural Experiment Station and by the DOE's BER, National Institute for Global Envi ronmental Change (NIGEC), Southeast Regional Center under Cooperative Agreement No. DE-FC03-90ER6101O. Ann Stodola, Craig Green, Jenny Moore, Xiangrong Duan, Brendon Johnson, Dr. Arnold Saxton, Dr. Richard M. Evans, and Joshua Olinick were instrumental in accomplishing the work. Support for G. Michael Gebre in the development of Chapter 8 was provided by an appointment to the Oak Ridge National Laboratory (ORNL) Postdoctoral Research Associates Program administered jointly by ORNL and the Oak Ridge Institute for Science and Education. The authors of Chapter 8 thank Donald E. Todd and Paul J. Hanson for the extensive effort involved in sample collection and Ramie V. Wilkerson for excellent technical support. The authors of Chapter 9 thank Paul J. Hanson, Michael Huston, Donald E. Todd, and Scott Dolvin for assistance in obtaining samples from the TDE site and for assistance with sample processing. We thank the staff at the Biology Station of Chamela, Universidad Nacional Aut6noma de Mexico for assistance with our work at Chamela. Sandy McLaughlin provided assistance with the tree core sectioning. The authors of Chapter 13 thank Patrice Cole, Caroline DeV an, Leigh Thomas, and Don Todd who helped collect, process, and plant seeds and assisted in the construction and maintenance of the seed-planting experiment. Paul Hanson en couraged our initiation of this project on the TDE, assisted in the development of our soil-water monitoring protocol, loaned us TDR probes and temperature dataloggers, and collated and summarized the temperature dataset. Travis Belote improved earlier drafts of the manuscript. Research on Chapter 13 was supported in part by the Department of Ecology and Evolutionary Biology, University of Tennessee, Knoxville. The authors of Chapter 14 thank the following individuals for their facilitation of the collection of field data: D. Boldt, Danim Moe, Nilong R. Parikh, Jorge Quiles, Shawnette Warner, and Donald E. Todd. The authors of Chapter 16 acknowledge the cooperation of the Public Power Institute of the Tennessee Valley Authority with Oak Ridge National Laboratory. They also thank Emily Childs, Brian Hale, Alan Mays, Jennifer Martin, Anita Rose, Larry Shelton, Brian Sunderland, Erich Webber, and numerous students for assistance in the collection and extraction of root samples and in the photograph ing and analyzing of minirhizotron data. Thanks are also due to E.G. O'Neill, Ron Hendricks, and Kurt Pregitzer for supplying minirhizotron equipment, assis tance, and advice. The authors of Chapter 18 thank Kenny Peavy, Larry Wilson, and Sue Car stensen for help in field sampling and/or laboratory analyses. George Cotsonis provided statistical advice and assistance in analyzing the data, and Chris Mowry assisted with the graphical presentation of the final results. Funding was provided by DOE's BER, NIGEC, Southeast Regional Center under Cooperative Agree ment No. DE-FC03-90ER6101O.