Physiological Ecology of. North American. Plant Communities. EDITED BY.
BRIAN F. CHABOT. Section of Ecology and Systematics. Cornell University. AND.
Physiological Ecology of North American Plant Communities
W. DWIGHT BILLINGS,
Wyoming, 1978. Photograph by Kim Peterson
Physiological Ecology of North American Plant Communities EDITED BY
BRIAN F. CHABOT Section of Ecology and Systematics Cornell University AND
HAROLD A. MOONEY Department of Biological Sciences Stanford University
NEW YORK
LONDON
CHAPMAN AND HALL
First published 1985 by Chapman and Hall 733 Third Avenue, New York NY 10017 Published in Great Britain by Chapman and Hall Ltd 11 New Fetter Lane, London EC4P 4EE © 1985 Chapman and Hall Softcover reprint of the hardcover 1st edition 1985
All rights reserved. No part of this book may be reprinted or reproduced or utilized in any form or by any electronic, mechanical or other means, now known or hereafter invented, including photocopying and recording, or in any information storage and retrieval system, without permission in writing from the publisher.
Ubrary of Congress Cataloging in Publication data Main entry under title: Physiological ecology of North American plant communities. Includes bibliographies and index. 1. Botany - North America - Ecology. 2. Plant communities North America. 3. Plant physiology. 1. Chabot. Brian F. II. Mooney. Harold A. QKllO.P49 1985 581.5'097 84-9586 ISBN-13: 978-94-010-8641-7 e-ISBN-13: 978-94-009-4830-3 DOl: 10.1007/978-94-009-4830-3 British Ubrary Cataloguing in Publication Data Physiological ecology of North American Plant communities. 1. Botany-North America-Ecology 1. Chabot, Brian F. II. Mooney. H.A. 581.5'097 QKllO
Contents
Preface List of contributors
1
The historical development of physiological plant ecology
xi xiii
1
W.D. BILLINGS
1.1 1.2 1.3 1.4 1.5 1.6
2
The beginnings The 19th century The spread of physiological ecology to North America The middle decades The post-war revival of physiological ecology Some thoughts about the future of physiological ecology References
Arctic
1 3 6 9 10 12 13
16
F. STUART CHAPIN III AND GAIUS R. SHAVER
2.1 2.2 2.3 2.4 2.5 2.6 2.7 2.8 2.9 2.10 2.11
Introduction Environment Vegetation and growth forms Phenology and growth Photosynthesis and respiration Water relations Mobile carbon pools Nutrient absorption, storage, and loss Allocation Reproduction Summary References
16 16 20 22 24 26 27 29 30 31 32 32
Contents
vi 3
Alpine
41
L.C. BLISS
3.1 3.2 3.3 3.4 3.5 3.6 3.7 3.8 3.9 3.10 3.11
4
Introduction Environment Floristics Plant communities and environmental gradients Seed germination and seedling establishment Growth forms and plant growth Plant phenology and growth Physiological responses Timberline The role of snow Summary References
Taiga
41 41 45 45 47 48 52 54 57 59 60 60 66
WALTER C. OECHEL AND WILLIAM T. LAWRENCE
4.1 4.2 4.3 4.4 4.5 4.6 4.7 4.8 4.9 4.10 4.11 4.12 4.13 4.14 4.15
Introduction Vegetation zones of the taiga Environment Growth forms and phenology Controls on carbon balance Temperature response Light response Water stress Nutrient relations Production Causes of timberline Fire ecology Establishment and reproduction Succession Summary and research needs References
5 Western montane forests
66 67 69 72
73 75 76 78 80 80 82 83 83 88 90 91 95
W.K. SMITH
5.1 5.2 5.3 5.4
Introduction Environment Ecophysiology Summary and perspectives References
95 98 101 114 120
Contents 6
Coniferous forests of the Pacific Northwest
vii 127
JAMES P. LASSOIE, THOMAS M. HINCKLEY AND CHARLES C. GRIER
6.1 6.2 6.3 6.4 6.5
Introduction Community structure Physiological characteristics and responses Tree structure Conclusion References
7 Annuals and perennials of warm deserts
127 128
l35
149 153 154
162
JAMES EHLERINGER
7.1 7.2 7.3 7.4 7.5 7.6
Introduction The physical environment Phenology and life cycle adaptations Leaf and canopy adaptations Photosynthetic and water relations adaptations Summary References
8 Desert succulents
162 162 164 167 170
177 177 181
PARK S. NOBEL
8.1 8.2 8.3 8.4 8.5
9
Introduction Water relations CO 2 uptake and acidity changes Thermal relations Conclusions References
Cold desert
181 182 185 190 193 195 198
M. CALDWELL
9.1 9.2 9.3 9.4 9.5 9.6 9.7
10
Introduction Photosynthesis in the cold desert environment The moisture constraint The salinity constraint Carbon balance of cold desert shrubs Nitrogen Summary: stress in the cold desert References
Chaparral
198 200 203 206 207 208 210 211 213
H.A. MOONEY AND P.C. MILLER
10.1 Extent and general character 10.2 Early ecological studies
213 213
Contents
viii 10.3 10.4 10.5 10.6 10.7 10.8 10.9
11
Environmental rhythms Growth forms and vegetation rhythms Water balance Carbon balance Nutrient balance Fire ecology Summary References
Grasslands
216 219 220 222 224 225 227 228 232
PAUL G. RISSER
12
11.1 Introduction 11.2 Plant response to environmental conditions 1l.3 Summary References
232 233 245 247
Deciduous forest
257
DAVID J. HICKS AND BRIAN F. CHABOT
12.1 12.2 12.3 12.4 12.5 12.6
13
Introduction Geography and vegetation Forest structure Plant response to seasonal environments Other periodic stresses Summary References
Tropical and subtropical forests
257 258 260 260 269 273 273 278
ROBERT W. PEARCY AND ROBERT H. ROBICHAUX
13.1 13.2 13.3 13.4 13.5
Introduction Distribution of tropical and subtropical forests Physiological ecology of tropical and subtropical forest species Physiological ecology of Hawaiian forest species Summary References
14 Marine beach and dune plant communities
278 278 279 281 292 293 296
MICHAEL G. BARBOUR, THEODORE M. DE JONG AND BR UCE M. PAVLIK
14.1 14.2 14.3 14.4 14.5 14.6
Introduction Light and temperature Water relations Salt spray Soil salinity Soil nutrients
296 299 304 306 307 308
Contents 14.7 Sand movement 14.8 Deductions from floristic analyses 14.9 Summary References
15 Coastal marshes
ix 312 314 316 318 323
B.L. HAINES AND E.L. DUNN
15.1 15.2 15.3 15.4 15.5 15.6
Index
Introduction Environmental stress Plant responses and adaptations Ecosystem properties affected by plant ecophysiology Management implications Conclusions and future research needs References
323 325 326 335 337 339 341 348
Preface
Although, as W.D. Billings notes in his chapter in this book. the development of physiological ecology can be traced back to the very beginnings of the study of ecology it is clear that the modern development of this field in North America is due in the large part to the efforts of Billings alone. The foundation that Billings laid in the late 1950s came from his own studies on deserts and subsequently arctic and alpine plants, and also from his enormous success in instilling enthusiasm for the field in the numerous students attracted to the plant ecology program at Duke University. Billings' own studies provided the model for subsequent work in this field. Physiological techniques. normally confined to the laboratory. were brought into the field to examine processes under natural environmental conditions. These field studies were accompanied by experiments under controlled conditions where the relative impact of various factors could be assessed and further where genetic as opposed to environmental influences could be separated. This blending of field and laboratory approaches promoted the design of experiments which were of direct relevance to understanding the distribution and abundance of plants in nature. Physiological mechanisms were studied and assessed in the context of the functioning of plants under natural conditions rather than as an end in itself. This approach was utilized by a large number of Billings' students as they moved out across North America, through his support and encouragement, to engage in research. They did not confine their activities to a single vegetation type, but rather as a group they covered most of the plant communities on the continent. It is because of this research breadth that the information was available to assemble this book. Not all ofthe authors of this book were students of Billings; however, they all felt his influence in one way or another. It is inevitable that certain plant communities received more attention than others as is evident from the book structure and contents. In particular, deserts and the arctic and alpine, have received a disproportionate share of concentrated studies. Important vegetation types, such as tropical forests, are just beginning to attract researchers to their varied problems.
xii
Preface
As noted earlier, knowledge from physiological ecology is necessary for understanding the patterning of plants on the landscape. Of equal importance, this knowledge is essential for a rational management of ecosystems. Studies from physiological ecology quantify the resource base of individual species, elucidate the specific mechanisms that have evolved to acquire and allocate these resources, and provide insights into the nature of species interactions. This knowledge can be utilized to specify the impact of various perturbations and stresses which are currently being imposed on ecosystems. Such information is currently being used to predict the effects of S02' acid rain, CO2 enhancement, etc. We know that species differ in their sensitivity to these factors. Knowledge of the precise differences can help us predict how species composition in a community might shift as the environment is modified or at what seasons or times species will be least susceptible to the impact of pollutants. In its modern configuration plant physiological ecology is an exciting developing field. The pace of its development has accelerated in recent years as sophisticated instrumentation has become available to quantify precisely the microenvironment of plants and their physiological responses in nature. The enormous diversity of the plant types which occur in North America provides the material for the development of a comprehensive comparative picture of the mechanisms which plants have evolved to cope with some of the hottest, coldest, wettest, and most arid habitats on earth. This book is an attempt to bring together some of the initial efforts in this field and it is our tribute to the leadership and inspiration of W.D. Billings.
Contributors
M.G. Barbour
Department of Botany, University of California, Davis, Davis CA 95616
W.D. Billings
Department of Botany, Duke University, Durham NC 27706
L.C. Bliss
Department of Botany, University of Washington, Seattle WA 98195
M.M. Caldwell
Department of Range Science, Utah State University, Logan UT 84322
B.F. Chabot
Section of Ecology and Systematics, Cornell University, Ithaca NY 14853
F.S. Chapin III
Institute of Arctic Biology, University of Alaska, Fairbanks AL 99701
T.M. DeJong
Department of Pomology, University of California, Davis, Davis CA 95616
E.L. Dunn
School of Biology, Georgia Institute of Technology, Atlanta GA 30602
J.R. Ehleringer
Department of Biology, University of Utah, Salt Lake City UT 84112
C.C. Grier
College of Forest Resources, University of Washington, Seattle WA 98195
B.L. Haines
Department of Botany, University of Georgia, Athens GA 30602
D.J. Hicks
Section of Ecology and Systematics, Cornell University, Ithaca NY 14853
xiv
Contributors
T.M. Hinckley
College of Forest Resources. University of Washington. Seattle WA 98195
J.P. Lassoie
Department of Natural Resources. Cornell University. Ithaca NY 14853
W.T. Lawrence
Centre for Energy and Environment Research. GPO Box 3682. San Juan. Puerto Rico 00936
P.C. Miller
Department of Biology. San Diego State University. San Diego CA 92182
H.A. Mooney
Department of Biological Sciences. Stanford University. Stanford CA 94305
P.S. Nobel
Department of Biology. University of California Los Angeles. Los Angeles CA 90024
W.C.Oechel
Systems Ecology Research Group. San Diego State University. San Diego CA 92182
B.M. Pavlik
Department of Botany. Iowa State University. Ames IA 50011
R.W. Pearcy
Department of Botany. University of California. Davis. Davis CA 95616
P.G. Risser
Illinois Natural History Survey. Natural Resources Laboratory. 607 East Peabody Drive. Champaign IL 61820
R.H. Robichaux
Department of Botany. University of California. Berkeley. Berkeley CA 94720
G.R. Shaver
The Ecosystems Center. Marine Biological Center. Woods Hole MA 02543
W.K. Smith
Department of Botany. University of Wyoming. Laramie WY 82071
