Vegetation Ecology PDF

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Vegetation Ecology A companion website with additional resources is available at www.wiley.com/go/vandermaarelfranklin/vegetationecology with Figures and Tables from the book Vegetation Ecology Second Edition Eddy van der Maarel & Janet Franklin University of Groningen, The Netherlands Arizona S...

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Vegetation Ecology

A companion website with additional resources is available at www.wiley.com/go/vandermaarelfranklin/vegetationecology with Figures and Tables from the book

Vegetation Ecology

Second Edition

Eddy van der Maarel & Janet Franklin University of Groningen, The Netherlands Arizona State University, USA

A John Wiley & Sons, Ltd., Publication

This edition first published 2013 © 2013 by John Wiley & Sons, Ltd. Blackwell Publishing was acquired by John Wiley & Sons in February 2007. Blackwell’s publishing program has been merged with Wiley’s global Scientific, Technical and Medical business to form Wiley-Blackwell. Registered office: John Wiley & Sons, Ltd, The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK Editorial offices: 9600 Garsington Road, Oxford, OX4 2DQ, UK The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK 111 River Street, Hoboken, NJ 07030-5774, USA For details of our global editorial offices, for customer services and for information about how to apply for permission to reuse the copyright material in this book please see our website at www.wiley.com/wiley-blackwell. The right of the author to be identified as the author of this work has been asserted in accordance with the UK Copyright, Designs and Patents Act 1988. 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, except as permitted by the UK Copyright, Designs and Patents Act 1988, without the prior permission of the publisher. Designations used by companies to distinguish their products are often claimed as trademarks. All brand names and product names used in this book are trade names, service marks, trademarks or registered trademarks of their respective owners. The publisher is not associated with any product or vendor mentioned in this book. This publication is designed to provide accurate and authoritative information in regard to the subject matter covered. It is sold on the understanding that the publisher is not engaged in rendering professional services. If professional advice or other expert assistance is required, the services of a competent professional should be sought. Library of Congress Cataloging-in-Publication Data Vegetation ecology / [edited by] Eddy van der Maarel & Janet Franklin. – 2nd ed. p. cm. Includes bibliographical references and index. ISBN 978-1-4443-3888-1 (cloth) – ISBN 978-1-4443-3889-8 (pbk.) 1. Plant ecology. 2. Plant communities. I. Maarel, E. van der. II. Franklin, Janet, 1959– QK901.V35 2013 581.7–dc23 2012018035 A catalogue record for this book is available from the British Library. Wiley also publishes its books in a variety of electronic formats. Some content that appears in print may not be available in electronic books. Front cover image: Vegetation mosaic in the calcium-poor coastal dunes in North Holland, with dune heathland, scrub, a dune lake and parabolic dunes in the background; photo Eddy van der Maarel (March 2005). Back cover image: Sonoran Desert scrub vegetation of the Arizona Uplands, also known as ‘saguaro-palo verde forest’, shown here in the Rincon Mountains, Saguaro National Park, east of Tucson, Arizona; photo Janet Franklin (April 2012). Cover Design by Design Deluxe Set in 10.5/12 pt Classical Garamond by Toppan Best-set Premedia Limited

1

2003

Contents

Contributors Preface

1

Vegetation Ecology: Historical Notes and Outline Eddy van der Maarel and Janet Franklin 1.1 1.2 1.3

2

xi xv

1

Vegetation ecology at the community level Internal organization of plant communities Structure and function in plant communities and ecosystems 1.4 Human impact on plant communities 1.5 Vegetation ecology at regional to global scales 1.6 Epilogue References

17 20 22 24 24

Classification of Natural and Semi-natural Vegetation

28

2.1 2.2 2.3 2.4 2.5 2.6 2.7 2.8 2.9 2.10

28 30 33 35 40 42 52 54 58 60

Robert K. Peet and David W. Roberts

Introduction Classification frameworks: history and function Components of vegetation classification Project planning and data acquisition Data preparation and integration Community entitation Cluster assessment Community characterization Community determination Classification integration

1 14

vi

3

Contents

2.11 Documentation 2.12 Future directions and challenges References

63 64 65

Vegetation and Environment: Discontinuities and Continuities

71

Mike P. Austin

3.1 Introduction 3.2 Early history 3.3 Development of numerical methods 3.4 Current theory: continuum and community 3.5 Current indirect ordination methods 3.6 Species distribution modelling or direct gradient analysis 3.7 Synthesis Acknowledgements References

4

Vegetation Dynamics

Steward T.A. Pickett, Mary L. Cadenasso and Scott J. Meiners 4.1 4.2 4.3

Introduction The causes of vegetation dynamics Succession in action: interaction of causes in different places 4.4 Common characteristics across successions 4.5 Summary Acknowledgements References

5

6

Clonality in the Plant Community

Brita M. Svensson, Håkan Rydin and Bengt Å. Carlsson

71 72 74 78 86 93 101 103 103

107 107 108 114 131 134 135 135

141

5.1 Modularity and clonality 5.2 Where do we find clonal plants? 5.3 Habitat exploitation by clonal growth 5.4 Transfer of resources and division of labour 5.5 Competition and co-existence in clonal plants 5.6 Clonality and herbivory Acknowledgements References

141 145 148 151 153 158 159 160

Seed Ecology and Assembly Rules in Plant Communities

164

Peter Poschlod, Mehdi Abedi, Maik Bartelheimer, Juliane Drobnik, Sergey Rosbakh and Arne Saatkamp 6.1 6.2

Ecological aspects of diaspore regeneration Brief historical review

164 166

Contents

6.3 6.4 6.5 6.6 6.7 6.8

Dispersal Soil seed bank persistence Germination and establishment Ecological databases on seed ecological traits Seed ecological spectra of plant communities Seed ecological traits as limiting factors for plant species occurrence and assembly 6.9 Seed ecological traits and species co-existence in plant communities References

7

Species Interactions Structuring Plant Communities Jelte van Andel 7.1 7.2 7.3 7.4 7.5 7.6 7.7 7.8

8

167 177 180 186 186 187 191 192

203

Introduction Types of interaction Competition Allelopathy Parasitism Facilitation Mutualism Complex species interactions affecting community structure 7.9 Assembly rules References

221 225 227

Terrestrial Plant-Herbivore Interactions: Integrating Across Multiple Determinants and Trophic Levels

233

Mahesh Sankaran and Samuel J. McNaughton

8.1 Herbivory: pattern and process 8.2 Coping with herbivory 8.3 The continuum from symbiotic to parasitic 8.4 Community level effects of herbivory 8.5 Integrating herbivory with ecosystem ecology References

9

vii

Interactions Between Higher Plants and Soil-dwelling Organisms Thomas W. Kuyper and Ron G.M. de Goede 9.1 9.2 9.3 9.4

Introduction Ecologically important biota in the rhizosphere The soil community as cause and consequence of plant community composition Specificity and selectivity

203 204 205 211 212 215 218

233 241 247 250 255 257

260 260 261 263 265

viii

Contents

9.5 9.6 9.7

Feedback mechanisms Soil communities and invasive plants Mutualistic root symbioses and nutrient partitioning in plant communities 9.8 Mycorrhizal networks counteracting plant competition? 9.9 Pathogenic soil organisms and nutrient dynamics 9.10 After description References

10

11

Vegetation and Ecosystem

Christoph Leuschner

275 278 279 279 281

285

10.1 The ecosystem concept 10.2 The nature of ecosystems 10.3 Energy flow and trophic structure 10.4 Biogeochemical cycles References

285 287 289 299 305

Diversity and Ecosystem Function

308

Jan Lepš 11.1 11.2 11.3 11.4 11.5 11.6

Introduction Measurement of species diversity Determinants of species diversity in the plant community Patterns of species richness along gradients Stability On the causal relationship between diversity and ecosystem functioning Acknowledgements References

12

268 274

Plant Functional Types and Traits at the Community, Ecosystem and World Level Andrew N. Gillison 12.1 12.2

The quest for a functional paradigm Form and function: evolution of the ‘functional’ concept in plant ecology 12.3 The development of functional typology 12.4 Plant strategies, trade-offs and functional types 12.5 The mass ratio hypothesis 12.6 Functional diversity and complexity 12.7 Moving to a trait-based ecology – response and effect traits 12.8 Plant functional types and traits as bioindicators 12.9 Environmental monitoring 12.10 Trait-based climate modelling

308 309 315 319 324 329 341 341

347 347 348 348 355 361 362 363 370 372 374

Contents

12.11 Scaling across community, ecosystem and world level 12.12 Discussion Acknowledgements References

13

Plant Invasions and Invasibility of Plant Communities

Marcel Rejmánek, David M. Richardson and Petr Pyšek 13.1 13.2 13.3 13.4 13.5 13.6 13.7

Introduction Definitions and major patterns Invasibility of plant communities Habitat compatibility Propagule pressure and residence time What are the attributes of successful invaders? Impact of invasive plants, justification and prospects of eradication projects References

14

Vegetation Conservation, Management and Restoration Jan P. Bakker 14.1 14.2 14.3

Introduction From agricultural exploitation to nature conservation Vegetation management in relation to a hierarchy of environmental processes 14.4 Laissez-faire and the wilderness concept 14.5 Management and restoration imply setting targets 14.6 Setting targets implies monitoring 14.7 Effects of management and restoration practices 14.8 Constraints in management and restoration 14.9 Strategies in management and restoration References

15

Vegetation Types and Their Broad-scale Distribution Elgene O. Box and Kazue Fujiwara

15.1 Introduction: vegetation and plant community 15.2 Form and function, in plants and vegetation 15.3 Vegetation types 15.4 Distribution of the main world vegetation types 15.5 Regional vegetation 15.6 Vegetation modelling and mapping at broad scales 15.7 Vegetation and global change References

ix 376 377 377 377

387 387 388 393 401 402 404 413 418

425 425 427 430 430 433 437 438 444 447 450

455 455 456 464 466 469 472 479 481

x

16

Contents

Mapping Vegetation from Landscape to Regional Scales Janet Franklin 16.1 16.2 16.3 16.4 16.5

Introduction Scale and vegetation mapping Data for vegetation mapping Methods for vegetation mapping Examples of recent vegetation maps illustrating their different uses 16.6 Dynamic vegetation mapping 16.7 Future of vegetation mapping research and practice Acknowledgements References

17

Vegetation Ecology and Global Change

Brian Huntley and Robert Baxter

486 486 489 490 495 500 501 502 503 503

509

17.1 Introduction 17.2 Vegetation and climatic change 17.3 Confounding effects of other aspects of global change 17.4 Conclusions References

509 510 518 525 527

Index

531

The color plate section can be found between pp. 272–273.

A companion website with additional resources is available at www.wiley.com/go/vandermaarelfranklin/vegetationecology with Figures and Tables from the book

Contributors

Mehdi Abedi, Institute of Botany, University of Regensburg, D -93040 Regensburg, Germany [email protected] Dr Mike P. Austin, CSIRO Ecosystem Sciences, GPO Box 1700, Canberra ACT 2601, Australia [email protected] Professor Dr Jan P. Bakker, Community and Conservation Ecology Group, University of Groningen, P.B. 14, NL-9750 AA Haren, The Netherlands [email protected] Dr Maik Bartelheimer, Institute of Botany, University of Regensburg, D -93040 Regensburg, Germany [email protected] Dr Robert Baxter, School of Biological and Biomedical Sciences, University of Durham, South Road, Durham DH1 3LE, UK [email protected] Professor Elgene O. Box, Geography Department, University of Georgia, Athens, Georgia 30602-2502, USA [email protected] Professor Mary L. Cadenasso, Department of Plant Sciences, University of California Davis, Mail Stop 1 1210 PES, One Shields Avenue, Davis, CA 956168780, USA [email protected]

xii

Contributors

Dr Bengt Å. Carlsson, Department of Ecology and Evolution, Uppsala University, Norbyvägen 18D, SE-752 36 Uppsala, Sweden [email protected] Dr Ron G.M. de Goede, Department of Soil Quality, Wageningen University, P.O. Box 47, 6700 AA Wageningen, The Netherlands [email protected] Dr Juliane Drobnik, Institute of Botany, University of Regensburg, D -93040 Regensburg, Germany [email protected] Professor Janet Franklin, School of Geographical Sciences and Urban Planning, Coor Hall, 975 S. Myrtle Ave., Fifth Floor, Arizona State University, P.O. Box 875302, Tempe AZ 85287-5302, USA [email protected] Professor Kazue Fujiwara, Laboratory of Vegetation Science, Yokohama National University, Tokiwadai 79-7, Hodogaya-ku, Yokohama 240-8501, Japan [email protected] Dr Andrew N. Gillison, Center for Biodiversity Management, P.O. Box 120, Yungaburra 4884 QLD, Australia [email protected] Professor Brian Huntley, School of Biological and Biomedical Sciences, University of Durham, South Road, Durham DH1 3LE, UK [email protected] Professor Dr Thomas W. Kuyper, Department of Soil Quality, Wageningen University, P.O. Box 47, 6700 AA Wageningen, The Netherlands [email protected] Professor Jan Lepš, Department of Botany, Faculty of Biological Sciences, University of South Bohemia, Branišovská 31, CZ-370 05 České Budějovice, Czech Republic [email protected] Professor Dr Christoph Leuschner, Plant Ecology, Albrecht-von-Haller-Institute for Plant Sciences, University of Göttingen, Untere Karspüle 2, D -37073 Göttingen, Germany [email protected] Professor Scott J. Meiners, Department of Biological Sciences, Eastern Illinois University, 600 Lincoln Avenue, Charleston, IL 61920-3099, USA [email protected]

Contributors

xiii

Professor Samuel J. McNaughton, Department of Biology, Syracuse University, 114 Life Sciences Complex, Syracuse, NY 13244-1220, USA [email protected] Professor Robert K. Peet, Biology Department, University of North Carolina, 413 Coker Hall, Chapel Hill, NC 27599-3280, USA [email protected] Dr Steward T.A. Pickett, Cary Institute of Ecosystem Studies, Box AB, Millbrook, NY 12545-0129, USA [email protected] Professor Dr Peter Poschlod, Institute of Botany, University of Regensburg, D -93040 Regensburg, Germany [email protected] Dr Petr Pyšek, Institute of Botany, Academy of Sciences of the Czech Republic, CZ-252 43 Průhonice, Czech Republic [email protected] Professor Marcel Rejmánek, Department of Evolution & Ecology, University of California Davis, 5337 Storer Hall, Davis, CA 95616, USA [email protected] Professor David M. Richardson, DST-NRF Centre for Invasion Biology, Stellenbosch University, Private Bag X1, Matieland 7602, South Africa [email protected] Professor David W. Roberts, Department of Ecology, Montana State University, Bozeman, MT 59717-3460, USA [email protected] Sergey Rosbakh, Institute of Botany, University of Regensburg, D -93040 Regensburg, Germany [email protected] Professor Håkan Rydin, Department of Ecology and Evolution, Uppsala University, Norbyvägen 18D, SE-752 36 Uppsala, Sweden [email protected] Dr Arne Saatkamp, Aix-Marseille Université – IMBE, Faculté des Sciences de St Jérôme, F-13397 Marseille cedex 20, France [email protected] Dr Mahesh Sankaran, National Centre for Biological Sciences, Tata Institute of Fundamental Research, GKVK, Bellary Road, Bangalore, India [email protected]; and

xiv

Contributors

Institute of Integrative & Comparative Biology, Faculty of Biological Sciences, 9.18 LC Miall Building, University of Leeds, Leeds LS2 9JT, UK [email protected] Professor Brita M. Svensson, Department of Ecology and Evolution, Uppsala University, Norbyvägen 18D, SE-752 36 Uppsala, Sweden [email protected] Professor Dr Jelte van Andel, Community and Conservation Ecology Group, University of Groningen, Centre for Life Sciences, P.O. Box 11103, 9700 CC Groningen, The Netherlands [email protected] Professor Dr Eddy van der Maarel, Community and Conservation Ecology Group, University of Groningen, Centre for Life Sciences, P.O. Box 11103, 9700 CC Groningen, The Netherlands Home address: De Stelling 6, 8391 MD Noordwolde fr, The Netherlands [email protected]

Preface

This book started as a multi-authored account on the many-sided topic of Vegetation Ecology (more commonly called plant community ecology in North America) because this modern field of science can hardly been treated by one or a few authors. In this second edition still more topics have been treated in separate chapters. As editors we have certainly had some influence on the choice and contents of the various chapters, but nevertheless the chapters are independent essays on important aspects of vegetation ecology. This edition consists of 16 chapters following an introductory chapter, three more than in the first edition. In addition to the 13 original chapters, which are all updated and adapted to the new structure (described below), we were able to include three new topical chapters. In connection with this new structure, the introduction no longer contains the mini-essays that were in the first chapter of the first edition. Instead we will refer to that chapter (van der Maarel 2005), as several authors in this book do, and present a simplified introduction in this edition. We have modified the sequence of topics by starting the this edition with the chapters (2–4) that deal mainly with the concept, structure, environmental relations and dynamics of plant communities. The second group of chapters (5–9) continue on the internal organization of plant communities. Subsequent chapters (10–12) deal with the structural and functional aspects and processes in plant communities as part of ecosystems. Here the emphasis is on the organization of plant communities in relation to the ecosystem of which they form a part. Chapters 13 and 14 deal with human impacts on plant communities in their ecosystem and landscape setting. The final chapters (15–17) address communities and geographically lar...