Ecological Studies Analysis and Synthesis Edited by W. D. Billings, Durham (USA) F. Golley, Athens (USA) O. L. Lange, Wiirzburg (FRG) J. S. Olson, Oak Ridge (USA) Volume 29 Vegetation and Production Ecology of an Alaskan Arctic Tundra Edited by Larry L. Tieszen With 217 Figures Springer-Verlag N ew York Heidelberg Berlin Library of Congress Cataloging in Publication Data Main entry under title: Vegetation and production ecology of the Alaskan arctic tundra. (Ecological studies; 29) Bibliography: p. Includes index. I. Tundra flora-Alaska-Barrow. 2. Primary productivity (Biology)-Alaska-Barrow. 3. Tundra ecology-Alaska-Barrow. 4. Primary productivity (Biology)-Arctic regions. I. Tieszen, Larry L. II. Series. QKI46.V43 581.5'264 78-14039 All rights reserved. No part of this book may be transiated or reproduced in any form without written permission from Springer-Verlag. © 1978 by Springer-Verlag New York Inc. Softcover reprint of the hardcover 1s t edition 1978 987654321 lSBN-13: 978-1-4612-6309-8 e-lSBN-13: 978-1-4612-6307-4 DOl: 10.1007/978-1-4612-6307-4 Preface This volume on botanical research in tundra represents the culmination of four years of intensive and integrated field research centered at Barrow, Alaska. The volume summarizes the most significant results and interpretations of the pri mary producer projects conducted in the U.S. IBP Tundra Biome Program (1970-1974). Original data reports are available from the authors and can serve as detailed references for interested tundra researchers. Also, the results of most projects have been published in numerous papers in various journals. The introduction provides a brief overview of other ecosystem components. The main body presents the results in three general sections. The summary chapter is an attempt to integrate ideas and information from the previous papers as well as extant literature. In addition, this chapter focuses attention on pro cesses of primary production which should receive increased emphasis. Although this book will not answer all immediate questions, it hopefully will enhance future understanding of the tundra, particularly as we have studied it in Northern Alaska. The synthesis of these primary producer papers as well as those of other ecosystem components is being published in "An Arctic Ecosystem: The Coastal Tundra at Barrow, Alaska" (J. Brown et aI., Eds.) and can be considered a companion volume. "Truelove Lowland, Devon Island, Canada: A High Arctic Ecosystem" (L. C. Bliss, Ed.) provides an invaluable comparison from the High Arctic, and "Fennoscandian Tundra Ecosystems" (F. E. Wielgolaski, Ed.) summarizes recent studies in northern Europe. Comparative information and the synthesis at the international level can be found in a number of recent books including: "Structure and Function of Tundra Ecosystems" (T. Rosswall and O. W. Heal, Eds.) and "Primary Production and Production Processes, Tundra Biome" (L. C. Bliss and F. E. Wielgolaski, Eds.). "Tundra; Compara tive Analysis of Ecosystems" (L. C. Bliss et aI., Eds.) is being prepared as an international synthesis. The rising environmental awareness in the world coincided with the discovery Preface VI of large oil deposits on the Alaskan Arctic Coastal Plain. The period 1970-1974 was therefore especially interesting as the U.S. Tundra Biome Program served as a scientific focus for the environmental interests of private industry, the U.S. Government and the public. In many ways we have all benefited from this confrontation and perhaps the resolution of future environmental issues will now be easier. The interest, encouragement, and support of these diverse groups is sincerely appreciated. Direct financial support of the program has been derived from three major sources: the National Science Foundation, the State of Alaska through the University of Alaska, and individual companies and members of the petroleum industry. The NSF funding was under the joint sponsorship of the U.S. Arctic Research Program (Office of Polar Programs) and the U.S. Interna tional Biological Programme (Ecosystem Analysis). The Office of Naval Research through its Naval Arctic Research Laboratory (NARL) at Barrow provided the field and laboratory support without which the U.S. Tundra Biome Program would not have been possible. Two former direc tors of NARL, Dr. Max C. Brewer and Mr. John Schindler, deserve particular credit for facilitating this support. Alpine field support was provided through the University of Alaska's Institute of Arctic Biology for Eagle Summit and through the University of Colorado's Institute of Arctic and Alpine Research at its Mountain Research Station for Niwot Ridge. Administratively, the Tundra Biome Center at the University of Alaska provided vital contractual and support services. Dr. George C. West, Director of the Tundra Biome Center, Dr. Keith Van Cleve, and Mr. David Witt deserve particular credit. The U.S. Army Cold Regions Research and Engineering Laboratory (CRREL) provided overall man agement, specialized logistic and equipment support and editing services to the program through an interagency agreement with the NSF and in the spirit of Public Law 40-280 which authorized the U.S. IBP. Mr. Stephen L. Bowen of CRREL and his staff, particularly Donna Murphy and Audrey White, were responsible for the preparation of all Biome reports and the final draft of this volume. Mr. Harold Larsen, CRREL illustrator, supervised the preparation of illustrations. In addition to the directly funded support, parent institutions, almost without exception, provided their staff members with a variety of on campus and other institutional aid. This is gratefully acknowledged at this time on behalf of the entire program. The success of this program is to a large extent due to the untiring optimism and perseverance of one person, Dr. Jerry Brown of CRREL, who served as director from 1970-1975. His managerial skills allowed many of us to concentrate more effort on scientific problems. Research in the U.S. Tundra Biome has been an extremely exciting and rewarding experience. To a large extent this has occurred because the program assembled more than 20 diverse projects most of which were committed to the "program," to full cooperation, and to an experimental assessment of plant function and productivity in tundra. It is perhaps this commitment to understand plant function and the basis for productivity which distinguishes our program. The dedication of most projects under the difficult conditions of remote field experimentation is also highly appreciated. Preface VII The program was enriched by the high degree of cooperation and criticism from international colleagues and U.S. participants in other biome programs. The opportunity for international meetings and exchanges as well as U.S. Specialists meetings is greatly appreciated for it helped clarify experimental approaches at regular intervals. May this international spirit of cooperation and camaraderie, especially among the young scientists, be one of the lasting results of the IBP Tundra Biome Program. L. L. TIESZEN References Bliss, L. C. (Ed.) (1977) Truelove Lowland, Devon Island, Canada: A High Arctic Ecosystem. Edmonton: University of Alberta Press, 714 p. Bliss, L. C. and F. E. Wielgolaski (Eds.) (1973) Primary Production and Production Processes, Tundra Biome. Proceedings of Conference, Dublin, Ireland, April 1973. International Tundra Biome Steering Committee, Stockholm, 256 p. Brown, J., P. C. Miller, L. L. Tieszen, F. L. Bunnell, and S. F. MacLean, Jr. (Eds.) (in prep.) An Arctic Ecosystem: The Coastal Tundra at Barrow, Alaska. Stroudsburg, Pa.: Dowden, Hutchinson and Ross. Rosswall, T. and O. W. Heal (Eds.) (1975) Structure and Function of Tundra Ecosystems. Swedish Natural Science Research Council, Ecol. Bull. NFR 20, 450 p. Wielgolaski, F. E. (Ed.) (1975) Fennoscandian Tundra Ecosystems, Parts 1 and 2. New York: Springer-Verlag, 366 and 337 p., respectively. Contents Preface v List of Contributors xvi Section I. Floristics, Vegetation, and Primary Production 1 la. Vascular Plants 1 lb. Nonvascular Plants 2 1. Introduction 3 LARRY L. TIESZEN Background 3 Research Design and Site Selections 5 Overview of the Barrow Ecological System 7 References 17 2. Vegetation, Floristics, and Phytogeography of Northern Alaska 19 D. F. MURRAY Tundra Landscapes and Vegetation 19 Floristics and Phytogeography 27 Acknowledgments 33 References 33 3. Spatial and Temporal Variation of the Vegetation and its Productivity 37 P.J. WEBBER Introduction and Site Overview 37 Methods 39 Results 43 Discussion 85 Summary and Conclusions 104 Acknowledgments 105 References 105 x Contents 4. Seasonal Dynamics of Above- and Belowground Production of Vascular Plants at Barrow, Alaska 113 J. G. DENNIS, L. L. TIESZEN and M. A. VETTER Introduction 113 Methods and Materials 115 Results 116 Discussion 134 Acknowledgments 138 References 138 5. Floristics, Phytogeography and Ecology of Arctic Alaskan Bryophytes 141 W. C. STEERE The Bryophyte Flora and its Floristic Elements 141 Ecology and Physical Factors of the Environment 149 Acknowledgments 163 References 164 6. Composition and Bryomass of the Moss Layers of Two Wet-Tundra- Meadow Communities near Barrow, Alaska 169 J. R. RASTORFER Introduction 169 Materials and Methods 169 Results 172 Discussion 177 Acknowledgments 181 References 181 7. The Role of Lichens in the Structure, Productivity, and Mineral Cycling of the Wet Coastal Alaskan Tundra 185 M. E. WILLIAMS, E. D. RUDOLPH, E. A. SCHOFIELD, and D. C. PRASHER Introduction 185 Environmental Setting 185 Lichen Occurrence 186 Methods 188 Results and Discussion 196 Conclusions 204 Acknowledgments 204 References 204 8. The Role of Algae in Tundra Soil 207 R. E. CAMERON, A. D. KNOX, and F. A. MORELLI Introduction 207 Methods 209 Results and Discussion 210 Conclusions 222 Contents xi Acknowledgments 223 References 224 9. Ecto- and Endomycorrhizae of Arctic Plants at Barrow, Alaska 229 O. K. MILLER, JR. and G. A. LAURSEN Introduction 229 Distribution of Ectomycorrhizae 232 Distribution of Endomycorrhizae 233 Distribution of Mycorrhizal Fungi 234 Carbon, Nitrogen, and Phosphorus 234 Conclusions and Discussion 236 Acknowledgments 236 References 236 Section II. Photosynthesis, Respiration and Water Relations. 239 IIa. Plant and Community Photosynthesis 239 lIb. Photosynthesis and Water Relations 240 10. Photosynthesis in the Principal Barrow, Alaska Species: A Summary of Field and Laboratory Responses 241 L. L. TIESZEN Introduction 241 Methods 242 Results and Discussion 242 Conclusions 265 Acknowledgments 266 References 266 11. Primary Production Processes in Arctic Bryophytes at Barrow, Alaska 269 W. C. OECHEL and B. SVEINBJORNSSON Introduction 269 Field Research Site 270 Methods 271 Results and Discussion 273 Conclusions 2% Acknowledgments 2% References 297 12. Meteorological Assessment of CO Exchange Over an Alaskan Arctic 2 Tundra 299 P. I. COYNE and J. J. KELLEY Introduction 299 Theory 299 Experimental Procedure 300 Data Analysis 301 3m ~wb xii Contents Discussion 314 Acknowledgments 318 References 319 13. Constraints on Tundra Productivity: Photosynthetic Capacity in Relation to Solar Radiation Utilization and Water Stress in Arctic and Alpine Tundras 323 M. M. CALDWELL, D. A. JOHNSON and M. FAREED Introduction 323 Methods 324 Results and Discussion 325 Acknowledgments 341 References 341 14. Some Aspects of Water Relations of Arctic and Alpine Regions 343 P. C. MILLER, W. A. STONER, and J. R. EHLERINGER Introduction 343 Conceptual Framework 344 Methods 345 Results 348 Discussion and Conclusions 353 Acknowledgments 356 References 356 15. Radio-Tracer Measurement of Transpiration in Tundra Vegetation, Barrow, Alaska 359 J. J. KORANDA, B. CLEGG, and M. STUART Introduction 359 Methods 359 Results and Discussion 361 Conclusions 367 Acknowledgments 368 References 368 16. Simulation of the Effect of the Tundra Vascular Plant Canopy on the Productivity of Four Moss Species 371 W. A. STONER, P. C. MILLER, and W. C. OECHEL Introduction 371 Simulation Models and Methods 371 Results and Discussion 375 Acknowledgments 385 References 385
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